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DOG COLOR
DOGICA® 3D World of Dog & Puppy

DOG COLOR















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Dog color
Dog Color Genotypes and Genetics
Dog and Puppy Fur and Nose Colors
Variety of Dog Eyes and Coat Colors
Extreme Dog Color Schemes
Free Dog Coloring Templates
Breeding Dogs in Colors
White vs Albino Dogs
Dog Nose Colors
Dog Eyes Colors
Albino Dogs


DOG COLORS

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Dog Color
DOG GENOTYPES & COLORS
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B/- D/- E/- K/- = black
b/b D/- E/- K/- = brown (chocolate)
B/- d/d E/- K/- = blue
b/b d/d E/- K/- = fawn
("-" is either the dominant or recessive allele)

AGOUTI:
at^at B/- D/- E/- k/k = black with tan points
at^at b/b D/- E/- k/k = chocolate with tan points
at^at B/- d/d E/- k/k = blue with dilute tan points
at^at b/b d/d E/- k/k = fawn with dilute tan points

NON-EXTENSION RED (cream):
B/B d/d e/e = dilute red to pale cream with gray nose (dog is genetically a dilute black, but will be a cream color)
B/b d/d e/e = dilute red to pale cream with gray nose (dog is genetically a dilute black, but will be a cream color)
b/b d/d e/e = dilute red to pale cream with rosey-brown nose (dog is genetically dilute brown, but will be cream color)
b/b D/d e/e = dilute red to pale cream with brown nose (dog is genetically brown, but will be cream color)
b/b D/D e/e = dilute red to pale cream with brown nose (dog is genetically brown, but will be cream color)
B/B D/D e/e = dilute red to pale cream with black nose (dog is genetically black, but will be cream color)
B/b D/d e/e = dilute red to pale cream with black nose (dog is genetically black, but will be cream color)








Color of Dog
GENE-RELATED
DEFINITIONS

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First of all, please aknowledge and get familiar with the following gene-related definitions:

Gene: Genes control just about everything about a living creature. They control how the creature grows and develops, how it functions and how it looks too. On this site we're focusing on just one aspect of looks - colour.A gene is basically a set of instructions which tells the cell how to produce a particular protein (in our case, pigment). The genetic make-up of a creature is described as its genotype.

Chromosome: The nuclear structure which houses (contains) the genetic information. Chromosomes exist in pairs and therefore there are always two copies of a given gene.

Locus \ Locci: the position of a gene on a chromosome. Every gene has a specific locus. Genes are all tied together into a long strand of DNA. Each point on this strand of DNA is called a locus (locii is the plural). At each locus there are two alleles forming the gene.
The alleles present at each locus are picked from a list of possible alleles, which is called a series. There may be any number of alleles in a series, but it's usually 2-5. Each locus has its own series, and the alleles in that series can only occur at that locus. For example, in the E series in dogs there are three alleles - E, Em and e. Each dog has a combination of two of those alleles from the E series on its E locus. It might have one copy of e and one of E, or maybe two copies of Em. See the "Summary of series" page for a list of all the series.

Genotype: the genetic make-up of an individual

Phenotype: While a genotype is the genetic makeup of an organism (a technical list of locii, genes and alleles), phenotype describes the effect of the genotype on the look of the organism. If we talk about a black dog with white markings then we are talking about its phenotype. If we talk about a KkSii dog then we are talking about its genotype.

Homozygous: the condition when both alleles of a gene pair are identical

Heterozygous: the condition when both alleles of a gene pair are different

Dominant: term describing a gene which can produce a phenotype when present only once; also the

Phenotype which results

Recessive: term describing a gene which must be present twice to produce a phenotype; also refers to the phenotype which results

Wild: the "normal" phenotype

Mutant: the non-normal phenotype; is a relative term (relative to the population from which the organism originates

Color Genes: genes that affect the pigment color of hairs

Pattern Genes: genes that affect the distribution of a particular color.

Alleles: Different variants of genes are known as alleles. Alleles come in pairs - one from each parent. When we write about alleles we use letters to denote them, for example a and b.








ALBINISM IN DOGS, ALBINO DOG
ALBINISM IN DOGS
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There are two main types of albinism:
Oculocutaneous - eyes, skin, and hair.
Ocular - just eyes.

There are also different levels of albinism, and different types within those two classifications. In fact, until recently, a lot of people denied they even existed. Sure, white dogs are everywhere!

But albino dogs aren't just white - they a complete or almost complete lack of pigment in the fur, skin, eyes, and nose. True albino dogs with no pigment at all are extremely rare.

ALBINISM IN DOGS, ALBINO DOG and PUPPY

In 1976, a white female Doberman Pinscher named Padula's Queen Shebah was born to two black and rust parents. Shebah was bred to her son to produce more white Dobermans. Since then, several thousand Dobermans, many of them white, have descended from Shebah. Although they are called white, they are actually light cream, with blue eyes and pink nose, lips and eye rims. The Doberman Pinscher Club of America (DPCA), backed by several geneticists, contended they were albinos. But many of their breeders insisted they weren't because they didn't fit the image of the prototypical albino with white fur and pink eyes.

ALBINISM IN DOGS, ALBINO DOG and PUPPY

Dobermans are the only breed in which albinos appear to be purposefully bred, very much against the wishes of the DPCA. They squint in bright light, and they're prone to sunburn, which may lead to skin tumors. A recent study (Winkler PA, Gornik KR, Ramsey DT, Dubielzig RR, Venta PJ, et al. (2014) found albino Dobes had a much higher incidence of eye and skin melanocytic tumors than normally pigmented Dobes.

ALBINISM IN DOGS, ALBINO DOG and PUPPY
What Causes Albinism?
A completely pure white animal with pink eyes and pale pink skin is considered albino, and is technically referred to as tyrosinase-negative. Tyrosinase is an enzyme involved in melanin production, and melanin is the natural substance that gives color (or pigment) to the eyes, skin, and hair. Albinism is a lack or defect of this enzyme and is caused by a recessive gene that is inherited from both parents. Many true albino dogs will suffer from deafness because the unpigmented skin in the ear canal causes the nerve endings to degenerate.

ALBINISM IN DOGS, ALBINO DOG and PUPPY

The truth is, more than 60 different gene mutations in various species are known to cause albinism, often with slightly different effects. Light fur with blue eyes is typical of the most common type of albinism, oculocutaneous albinism type 2 (OCA2), in humans. It is now acknowledged that white Dobermans are in fact albinos, and the causative gene has recently been discovered.

The mutation is not a part of any of the known dog color loci. This includes the C series, where mutations causing albinism in many others species, including cats, rats and mice, have been identified. However, mutations in SLC45A2 cause the OCA4 type of albinism responsible for cream-colored Bengal tigers, horses, and gorillas, as well as some albino humans.

ALBINISM IN DOGS, ALBINO DOG and PUPPY

But Dobes are not the only breed with albinos. Pekingese seem to have the second largest number, but they have also been seen in Shih Tzu, Poodles, Pit Bulls, Beagles, Pugs, Dachshunds and doubtless, many others. In the few individuals tested, they don't seem to share the same mutant gene with the albino Dobes, nor do they seem to share the C-series allele seen in so many other domestic albino animals. In no breed is the purposeful breeding of these dogs encouraged. Nonetheless, if you have one, they make fine companions but you need to take steps to keep them from being dazzled by bright light or from getting sunburned.

ALBINISM IN DOGS, ALBINO DOG and PUPPY

Despite the fact that albinism is equally rare among all vertebrates from humans to fish and birds, the real albino dogs are even rarer due to two reasons. The first reason is that the most "albino" dogs pictures on the internet depicting pretty normal dogs with white fur. The second one, it's a bit hard to figure out if your dog is real albino.









WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY
WHITE vs ALBINO DOGS
PARTIAL ALBINISM IN DOGS

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HOW TO DISTINGUISH
WHITE vs ALBINO DOG ?

Tip #1 - Inspect the dog's coat. If the dog is predominantly white, but has some brown or black fur around the nose or feet, the dog is not an albino. The albino dog is unable to produce hair color of any kind.

Tip #2 - Spread the fur and look at the dog's skin. Many dogs have mottled skin with large colored patches beneath their fur. These are normal skin colorations and prove that the dog has the ability to produce melanin, the key component in skin coloration. Albinos lack the ability to produce melanin.

Tip #3 - Look at the dog's eyes. If the iris of the eyes is brown, the dog has normal melanin distribution. If the iris of the eyes is pink, the dog is an albino. The iris appears pink because the eyes lack the pigment to shelter the blood vessels of the eye.

WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY

Albinism is rare in all animals, including dogs, and many people easily confuse white-coated dogs, or dogs that exhibit forms of albinism, for albinos. True albinism is a genetic condition in which pigmentation of eyes, coat, and skin is completely absent. An important distinction to draw between dogs with white coats and albino dogs is that white-coated dogs produce the color white, while albinos only appear white due to lack of pigmentation.

WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY

All-white dogs have genetic markers by virtue of which the white pigment masks, or suppresses, all other colors. Albinism, on the other hand, is caused by the absence of both melanin and the enzymes that produce it. That said, some dogs exhibit characteristics of albinism without being true albinos. Let's explore the distinctions, as plainly and legibly as possible, and see what makes an albino dog an albino.

WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY
Partial albinism in dogs
Some dogs may appear to be true albinos, but retain some pigmentation, which will be most noticeable on the nose or stomach. We can call this partial albinism, but there is actually a range of melanins, and as such, a wide variety of albinisms are possible and observable in dogs. Eyes and skin of albino dogs may appear pink, but it is the diffused color of blood vessels:

WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY

In cases of partial albinism, dogs produce only a small amount of melanin, sufficient to produce limited coloration. With the exception of small areas of pigmentation, whether in eyes, skin, or coat, what remains will retain that extremely pale, color-drained appearance.


WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY
Coat patterns confused with albinism
Instances of limited coloration in non-albino dogs produces two coat patterns, each producing limited color swatches on a dog's coat and skin. These patterns are known in breed standards and kennel clubs as "piebald" and "merle." Piebald dogs have mostly white-colored coats that display large spots or patches of dark coloration. Merle-coated dogs exhibit splotches or patches of color, not only on the coat, but on the skin as well.

WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY

Dogs with merle coats are also prone to having heterochromatic, or different-colored, eyes. As in white cats, the genes responsible for coat color, eye, and ear health are not causally linked, so white and albino dogs are not necessarily more likely to be born blind or deaf. However, a rare genetic combination, known as "double merle", does carry inherent health risks. Double merle dogs, like Keller in the photo above, may be mistaken for albino dogs. Unlike true albino dogs, who, aside from light sensitivity, are generally healthy, double-merle-coated dogs are at higher risk for both deafness and blindness. This beautiful dog is Keller, a double merle Australian Shepherd. Her owner writes very movingly about the difficulties and health issues of double merle dogs, not to be confused with albinos:

WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY
Light sensitivity in albino dogs
Melanin serves a number of uses in the body aside from providing pigmentation. In the eyes, the presence of melanin is one thing that allows dogs to process and filter light. For a true albino dog, without melanin or without much, direct sunlight causes pain in their eyes which makes them squint. True albino dogs should get minimal and carefully managed exposure to direct sunlight.

WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY

Light filtration is not the only purpose for melanin. With regard to the skin and body, it provides natural protection from the sun, as well as contributing to the body's ability to fight off infection. Further, albino dogs are far more prone to sunburn and to developing skin cancers due to their extreme photosensitivity.


WHITE vs ALBINO DOGS, ALBINISM IN DOGS, ALBINO DOG and PUPPY

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dog and puppy colors
CONTENTS OF LOCUSes
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A locus
("agouti series", affects distribution of both eumelanin and phaeomelanin)


dog and puppy colors dog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colors dog and puppy colors

- Ay: Sable (red with or without black tipping). Used to be labelled ay when dominant black was believed to be above it on the A locus, but now black has been given its own locus (K) and sable has been promoted to the most dominant on its locus (so has gained a capital letter!).
- aw: Agouti (banded hairs). Like sable above, but the hairs all over are banded with black. This is most likely the gene responsible for wolf grey.
- at: Tan points (black body with red on muzzle, chest, eyebrows, legs and vent). Only dominant over recessive black (below), so a dog needs to be homozygous for tan points in order to express them (or heterozygous for recessive black and tan points, but recessive black is very rare).
- a - Recessive black (solid black with no red in the coat at all).






B locus
("liver series", affects colour of eumelanin)


dog and puppy colorsdog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colorsdog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colorsdog and puppy colors dog and puppy colors

- B: Normal pigment. A Bb or BB dog produces normal black eumelanin.
- b: Liver pigment. A bb dog produces liver eumelanin instead of black.



D locus
("dilution series", affects intensity of eumelanin)


dog and puppy colorsdog and puppy colorsdog and puppy colors dog and puppy colorsdog and puppy colorsdog and puppy colors

Dillute Fake

dog and puppy colorsdog and puppy colorsdog and puppy colorsdog and puppy colorsdog and puppy colors

- D: Normal pigment.
- d: Diluted pigment. When homozygous (dd), turns black to blue and liver to isabella.

Breeds Carrying Dilution
American Staffordshire Terrier
Great Dane
Pomeranian
Beagle
Greyhound
Rat Terrier
Bearded Collie
Harrier
Russian Toy Terrier
Border Collie
Italian Greyhound
Schipperke
Borzoi
Kelpie
Shar Pei
Canary Dog (Presa Canario)
Miniature Pinscher
Shih Tzu
Cane Corso
Mudi
Slovakian Pointer
Chesapeake Bay Retriever
Neapolitan Mastiff
Staffordshire Bull Terrier
Chihuahua
Newfoundland
Thai Ridgeback
Chow Chow
Otterhound
Tibetan Mastiff
Dobermann
Peruvian Inca Orchid
Weimaraner
Finnish Lapphund
Plott Hound
Whippet
Foxhound (American and English)
Podengo Portugueso
Xoloitzcuintle
German Pinscher




E locus
("extension series", affects distribution of eumelanin)


DOG and PUPPY COLOR

DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR DOG and PUPPY COLOR

Recessive Red

DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR DOG and PUPPY COLOR

Grizzle & Domino

DOG and PUPPY COLORDOG and PUPPY COLOR

- Em: Masked (black on the muzzle and the ears, and sometimes spreading to black tipping on the chest and/or back).
- E: Normal extension (no restriction of pigment). Usually written with a capital letter, but is in fact recessive to Em.
- Eg: Grizzle/domino (found only in Salukis and Afghan Hounds)
- e: Recessive red (solid red all over, except for white markings). An ee dog is unable to produce any eumelanin (black) in its coat. Any black present will be turned to red. The eyes and nose are, however, unaffected (a recessive red dog may still have a black nose).






G locus
("greying series", affects how eumelanin keeps its itensity over time)


DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR

Premature Greying

DOG and PUPPY COLORDOG and PUPPY COLOR

- G:
Progressive greying. A dog with one or two G genes will be born dark-coloured and its hair will lighten over time.
- g: Normal (no lightening of pigment).

Breeds Carrying Greying Gene
Old English Sheepdog
Bearded Collie
Tibetan Terrier
Polish Lowland Sheepdog
Bedlington Terrier
Dandie Dinmont Terrier
Basset Griffon Vendeen (Grand and Petit)
Poodle (Standard, Miniature and Toy)
Irish Wolfhound
Deerhound
Cesky Terrier
Kerry Blue Terrier
Havanese
Glen of Imaal Terrier




H locus
("harlequin series", modifies merle)


DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR

Typical harlequin (HhMm)

DOG and PUPPY COLOR

Harlequin with the dilution gene, turning the patches and nose blue (ddHhMm)

DOG and PUPPY COLOR

Harlequin with the sable and masking genes (AyAyEmEmHhMm)

DOG and PUPPY COLOR

Harlequin with the brindle gene (sable is also necessary for the brindle to show all over the body) (ayaykbrkbrHhMm). The brindle will be broken up and patchy because of the merle gene.

DOG and PUPPY COLOR

Harlequin with irish spotting (HhMmsisi). The red line shows the border between the harlequin markings and the white spotting.

DOG and PUPPY COLOR

Merle without the harlequin gene (hhMm)

DOG and PUPPY COLOR

Lightly-marked harlequin, most likely a double merle (HhMM)

DOG and PUPPY COLOR

- H: Harlequin. Areas between patches on a merle dog are turned to white, leaving solid pigmented patches on a white base. Only works with merle gene and does not affect non-merle dogs.

- h: Non-harlequin (normal expression of merle).

DOG and PUPPY COLOR

I locus
("Intensity series", affects the richness of phaeomelanin)


DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR

Urajiro

DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR DOG and PUPPY COLOR

Husky Color

DOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLORDOG and PUPPY COLOR

Schnauzers

DOG and PUPPY COLORDOG and PUPPY COLOR

Dilution of phaeomelanin in dogs is now thought to be due to the Intensity locus. Currently, it is not known how this locus works or which alleles are present on it. However, the general idea is that is causes the phaeomelanin in the coat to lighten or darken. The above photos show the range of phaeomelanin colour in recessive red and sable dogs. The I locus does not affect eumelanin in any way, and so doesn't affect the nose or the eyes. The "dudley" noses and unpigmented eye rims sometimes seen in dogs with light-coloured phaeomelanin are in fact generally due to the recessive red gene and not the phaeomelanin dilution itself.

- alleles not yet known




K locus
("black series", "Brindle" affects eumelanin)





Variations In Base Color

The base colour on a brindle can vary as much as red can usually vary, from a deep Irish Setter colour to a light cream.
A light base coat on a brindle is generally due to the Intensity locus, which dilutes phaeomelanin (red pigment) to cream, but has no effect on eumelanin (black pigment).
These are all details taken from photographs of brindles with black stripes.

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Brindle with a Mask

A dog which has at least one Em gene on its E locus will have a black mask. This is an area of eumelanin covering the muzzle and sometimes the ears. If a dog is a brindled sable (brindle gene + sable gene), it'll be brindle all over, and on dogs like this masks may be visible.
You may notice that on the Great Dane above the phaeomelanin (red) is deeper on the back and on the chest and legs is almost white. This is most likely due to the urajiro pattern (see the C and I locus page). As urajiro only affects phaeomelanin, the black stripes are unaffected.

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Silver Brindle

Sometimes a brindle dog with black pigment may have silver stripes, usually on a cream background (never deep red). This colouration is generally associated with sighthounds, particularly Afghan Hounds. The overall effect may be similar to how a blue brindle (dd dilution gene) would look. Silver brindle is caused by the greying gene (G locus), which turns black to grey as the dog ages.
Sometimes dogs with black stripes and a very light cream base are also referred to as silver brindles.

dog and puppy colors

Brindle and Long Fur

Long and wirey fur can obscure brindle markings by making the stripes less distinct.
Even between longhaired breeds, the visibility of brindle can vary. The Dachshund here has very distinct stripes and is obviously a brindle, but the Skye Terrier just appears to be a muddy grey/brown. The main way we can tell this Skye is brindled is simply its muddy-coloured coat - this colour occurs almost exclusively on brindles and never on normal solid reds or sables. It also appears to have some lighter and darker areas, which also suggest it is a brindle.
This Glen of Imaal's brindling is very obvious on the short fur on its body, but less obvious on the longer fur on its head. If the Skye above were to be shaved down, it would look something like this.

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The Wolfhound and Deerhound show brindling on wirey coats. On the first two dogs, greying has caused the black striping to fade out.

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Brindle Tan Points and Saddle Markings

Because the brindle gene allows expression of the A locus, any A locus coat patterns can show brindling on their red sections. The main patterns are sable (AyAy) and tan points (atat). A sable brindle will appear solid brindle (in fact, all solid brindles are sables), although any areas of shading will remain dark. A tan pointed dog with the brindle gene will appear black with brindle points instead of tan.
All of these dogs display black-pigmented dogs with brindled points, with or without white markings.

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These two dogs are brindle with saddle markings(atat plus an unknown modifier, see the Agouti page). Note the large areas of solid black on the back and brindled tan on the legs and head. The Cardigan Welsh Corgi also shows white in the irish spotting pattern.

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Liver and Dilute Brindles

A brindle dog will have liver (brown) stripes if it is bb on the B locus, blue stripes if it is dd on the D locus, and isabella stripes if it is both dd and bb. Dilution affects phaeomelanin (red) too to some extent, so the base colour will be relatively light (certainly not a deep red colour, but more likely cream).
Here are three blue brindles. The Whippet is a very light brindle, and the blue colouration is mostly visible on the mask and nose. The Neapolitan Mastiff, on the other hand, is a heavy brindle, so the overall appearance is blue.

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"Missing" Stripes

On some light brindles there may be solid red areas where it appears stripes are "missing". This is particularly common in Whippets, but the genetic basis is unknown. Have a look at the Whippet below. In the first picture the brindle appears normal - however in the third picture you can see a large clear area on the dog's side.

Dog Colors Colorific Dog

Brindle Merles

Please, check the MERLES section.

- K: Black (solid black all over). Overrides A (agouti) series. Any genes on the A locus will not be expressed.
- kbr:Brindle (black stripes on a red base). kbr is dominant over k, so a dog only needs one kbr in order to be brindle (but will be overriden by one K gene). Brindle dogs will express whichever genes are on their A locus, but the red parts of the coat (phaeomelanin) will be brindled (black parts will not be affected).
- k: Non-solid black. A kk dog will express whichever genes are on its A locus.

Dog Colors








M locus
("merle series", affects intensity of eumelanin)


Blue Merles

The first dog has very few black patches, and they're mainly quite small. The second dog shows the normal merle pattern - a mixture of larger and smaller patches, covering roughly 50% of the body. This pattern is generally the most preferred in breed standards. The third dog has very large black patches, sometimes referred to as blanketing. The last dog is known as a minimal merle. It is almost completely black with just a very small amount of merling. This pattern is rare and generally discouraged because it can "hide" the merle gene if the black covers up all the merle in the coat. Dogs with little or no visible merling are sometimes called cryptic merles.

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The dogs above are called "blue merles" because of the bluish colour between the patches in their coat. This is a widely-used term but is actually misleading. Technically they should be "black merles". Their nose pigment is black and their eyes are brown or blue. They are able to make normal eumelanin in their coat, so their patches are black. If they didn't have the merle gene, they would be solid black. "Blue merle" is misleading because it seems to say that these dogs have blue pigment (dd acting on black - see Dilution page), when in fact they have black.

An example of the genotype of one of the above dogs would be: BBDDEEggMmSSKK (most of these aren't necessarily homozygous, but I will assume they are for ease, otherwise I'd have to keep saying, "KK, kK or Kk" etc). The genotype translates as: no liver colour (BB), no dilution (DD), no mask or recessive red (EE), no greying (gg), merle (Mm), no white spotting (SS), and solid black (KK).

Two solid blue merles (the Border Collie also has white in the irish spotting pattern and bronzing on its side).

This mixed breed could be classed as a cryptic merle as one side of the dog is almost completely solid black. The merling is only visible on the right side (first photo). This dog also has tan points (atat) and white spotting in the irish pattern.

Color of dogDOG COLOR

Examples of the variation in base colour of merles. In some cases the black may be diluted almost to white, but it is mostly somewhere between light "powder" grey and dark "grizzled" grey. It is not certain whether this variation has a genetic basis.

DOG COLOR DOG COLOR

Eyes and Noses

DOG COLOR

The random coat dilution caused by merle also affects the eyes and nose. The eyes may be all or partly blue, and the nose may be all or partly pink. Above are examples of "butterfly" noses on merles, which are partly pink.

The harlequin Great Dane here also shows a butterfly nose and wall eyes (one blue, one brown). Harlequin is a modified type of merle where the areas between the patches are diluted completely to white. See the Harlequin page for more information.

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Dilute Spots

Sometimes merles have patches that are only partially diluted, and are between the base and the patch colour. These are known as dilute spots, and they may sometimes appear brownish. The Australian Shepherd above has a large dilute spot on its knee.

DOG COLOR

Liver Merles



A normal "blue" merle becomes a "red" merle when it has bb on the B locus, i.e. when it has the liver gene. "Red merle" is also a misleading term because "red" is usually used to refer to phaeomelanin (tan, gold, cream etc) rather than eumelanin. A red merle should correctly be called a liver merle.
Liver turns all of the patches on a blue merle into brown and the colour between the patches becomes pale brown. As with all livers, liver merles have liver noses and amber eyes.
Here we have two long-haired Dachshunds. The dog at the back is a red (liver) merle with tan points, and the dog in the front is a normal blue (black) merle with tan points.

More examples of liver merles with tan markings and with or without white.

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Tan Markings

DOG COLOR

As the examples in the sections above have shown, merles can come with or without any of the tan patterns (A locus). The most common is normal tan points, but creeping tan and saddle patterns also occur.
All dogs which show tan markings must be kk on the K locus (K is dominant black, so if a dog has even one copy of it then they will be solid black, or solid merle if they are also Mm, regardless of what they have on the A locus). A tan pointed dog will be atat on the A locus (tan points are recessive and only dominant over recessive black, a, so a dog must have two at genes to display tan points).

The Norwegian Dunker is one of the only breeds where the saddle pattern and the merle gene occur together. This dog's pattern is between creeping tan and saddle. You can see that the merling covers the back, the tail, the back of the neck and the top of the head. The tan is very light and so is probably affected by the Intensity, which lightens red. This dog also has white in the irish spotting pattern. The Dunker photo was taken by June.



Masks



A dog which is Eme, EmE or EmEm on the E locus will have a mask (Em, the mask gene, is dominant). The mask can be black, liver, blue or isabella depending on the dog's pigment. If the dog also has the merle gene, the mask will be merled. Because of this, masks are not be visible on solid merle dogs - they're just merled like the rest of the coat. However, tan-pointed merles with little to no face white sometimes display masks (recognisable because the areas which should be tan on the muzzle are merled instead). Sable (ayay) dogs have masks which are much easier to see. A dog which is ayayEmEmMm (homozygous for sable, homozygous for mask, heterozygous for merle) will appear solid red with a merled mask.

Sable



A dog with AyAy on the A locus and kk on the K locus (allowing for the A locus markings to be visible) will be a sable. Sables are solid red with or without black tipping. Sometimes, especially if sable is combined with a mask, the black tipping can be quite extensive and can cause large black areas on the back, head and tail. In a sable merle, all of these black areas are merled. As you can imagine, how obvious it is that a dog is a sable merle depends heavily on how much black tipping they have. A sable merle with no black tipping (a "clear sable") will just appear to be a solid sable, possibly with one or two blue eyes (the only indication that they are a merle). In some breeds, such as Shetland Sheepdogs and Rough Collies, a slightly different version of sable exists. This type consists of brownish hairs on the back and head (even though these dogs have black pigment), and is often called "shaded sable". On this type of sable coat, merling can be quite visible (if there is a lot of dark brown shading) or very hard to see (if the shading is lighter and not so extensive). However, the merling is usually visible at birth, so breeders will generally know if their dogs are sable merles or just sables. A sable merle will have some faint, darker brown/tan patches on a lighter base, and the merling will usually be confined to the back and head. It is often most visible on the ears, where the fur is shorter.



Simon is also a shaded sable merle. These pictures show him from five months to 2 years old. His shorter puppy coat displays the merle much more clearly than his longer adult coat.

DOG and PUPPY COLOR DOG and PUPPY COLORDOG and PUPPY COLOR

Grizzle / Agouti

Haiku shows beautifully the effect of merle on agouti (aw). The agouti pattern consists of banded hairs scattered over the dog (and mostly concentrated on the head and back). As only the black (eumelanin) parts are affected by the merle gene, the overall effect of merle on grizzle is of scattered small black spots and flecks, with some patterns forming where the black would usually be on the grizzle (as can easily be seen in the photo of Haiku to the front, where the spots on his head form lines). These two alleles do not generally occur within the same breeds, so grizzle/agouti merle is very rare and is usually seen on Husky crosses. Haiku the Husky mix, photos by Dariana Kamenova:

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Brindle

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If a dog is kbrkbr (or kbrk, because kbr is dominant over k) on the K locus then any phaeomelanin (red) in their coat will be brindled. This applies to merle dogs too - there is no reason why a merle cannot also have brindle. If a merle dog is kbrkbr and has atat on the A locus then it will be merle with brindled tan points. If a merle dog is kbrkbr and AyAy (sable), it will be solid red with brindling all over ( and possibly with larger areas of black than is normal for a brindle on the back and/or head, if the sable has dark shading). The brindle in both the brindle-pointed merle and the solid brindle dog will be broken up into uneven spots rather than being complete stripes. This is because the stripes on a brindle are eumelanin (affected by merle, so random parts are diluted) and the base is phaeomelanin (not affected by merle).
A solid brindle merle is relatively easy to identify because of its broken-up stripes. However, when a merle dog has just brindle points it can be harder to recognise because the stripes are even less visible. The points will generally have a few black spots on them and will appear a darkish, muddy brown.

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The Cardigan Welsh Corgi is one of the few breeds where brindle and merle regularly occur together (the other main one being the Catahoula Leopard Dog). The dog on the left has tan points, but they are very dull and brownish compared to normal tan points (see the dog on the right, which does not have the brindle allele). This points to this dog being a brindle-pointed blue merle, which is very common in the breed. The brindle appears only on the points, where it is broken up by the merle gene. In this case, the dog is a very light-marked merle, so the points are very washed out with few spots. Brindle can also appear on merles with the creeping tan and saddle patterns.

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Ticking

DOG and PUPPY COLOR

A dog with TT or Tt on the T locus will have ticking on any white areas in their coat. It is thought that TT produces roan or heavy ticking and Tt produces lighter ticking. Ticking will only show up on a merle if it has white markings, and the ticking will be a mix of the merle base colour and the patch colour (so in a blue merle, black and grey).

Recessive Red

Recessive red is when a dog has the genotype ee on the E locus. This means it is unable to produce eumelanin (except in its nose and eyes), and can only produce phaeomelanin, so its coat is red all over (except for any white markings). Merle only affects eumelanin, so if there's none there, there can't be any merle. Just like with clear sable, a recessive red merle can be impossible to distinguish from a non-merle dog. It will simply appear solid red, and the only giveaway is if it has one or two blue eyes.

Dilution

Lastly, we need to mention dilutes. A dilute is a dog which is dd on the D locus. dd stops a dog from producing full eumelanin pigment and also affects phaeomelanin to a lesser extent, meaning that the pigment it produces is weak and pale. It turns a black dog into a blue dog with a blue nose and amber eyes) and a liver dog into an isabella. A merle with the dilution gene will appear very washed-out. The black or liver patches will be diluted almost to the same shade as the base colour. So a blue merle (a proper blue merle with the dilution gene, not a black merle like the ones we've been dealing with so far! will appear almost completely light grey with some faint darker grey patches, and an isabella merle will be light greyish brown with faint darker brown patches. On a longhaired dog, it can be next to impossible to tell a blue or isabella merle from a non-merle. To distinguish between a merle with dilution and a merle with greying (which produces much the same effect on the patches in the coat), look at the nose. If the dog has a blue nose, it has the dilution gene.

Double Merles

A double merle is one which is homozygous. That means it has two copies of the dominant M gene, so its genotype is MM. This impairs its ability to make pigment.
Double merles are typically mostly white and have merle patches on the head and sometimes the body. They do not necessarily have the allele for piebald, although if they do, they are likely to have even more white than double merles without a white spotting allele), but the MM genotype causes loss of pigment to such an extent that it tends to remain only on the upper areas of the dog - the top of the head, the back and the base of the tail, similar to the extreme spotting pattern. Sometimes double merles are even born completely white. The nose is generally mostly or all pink, and the eyes are generally wall or blue.



This Dachshund is a blue (black) double merle with tan called a double "dapple" in this breed). She has a large amount of colour on her body, probably because she has been bred from solid merle parents rather than merles with white spotting the amount of white on the merle parents tends to affect the amount of white on the double merle puppy, so high-white parents will generally produce higher-white puppies, but she does have a lot of white on her face. The eye on the side with the white is blue because of the lack of pigment, and she has a butterfly nose.



This Border Collie type has a typical double merle coat pattern. However, the full pigment on the nose and around the eyes suggests that she may in fact just be a high-white normal merle. It would be unusual for a double merle to have so much pigment.

Photos by Cat of Dog Rad Design



Here we have a probable harlequin double merle. Harlequins are merles with a modifier that changes the way the merle looks (see the page on merle modifiers). The same principles therefore apply to harlequin double merles as do to normal double merles - the patches will generally be smaller and confined to the head and the back, and there will often be a more pronounced lack of pigment around the eyes and nose.



The breeding of double merles is generally frowned upon, not just because double merle dogs rarely match breed standards (for example, a double merle Dachshund like Casey above would be disqualified for too much white and bad pigment), but also because of the health problems linked to them. Some double merles are perfectly healthy, but eye defects and deafness (due to lack of pigment in parts of the inner ear) are common, as well as sunburn and skin cancer on exposed pink areas.

Merle puppies are circled in blue and double merles are circled in red. Non merles are not circled.

It's clear from this that breeding a double merle to a non merle is the only way to get a completely merle litter. Double merle to non merle and merle to non merle are the only "safe" breedings - i.e. the only ones which cannot produce double merle puppies. Breeding merles of any sort (single or double) together is always risky. I haven't included a Punnet square for a double/double breeding, but that one is easy enough to work out - each parent can only possibly pass on the M gene, so all puppies will be MM (double). Because of this, double merles should never be bred together.









"Lethal White"

Double merles are often referred to as "lethal whites". This is an incorrect and misleading term. There is nothing lethal about the merle gene, it can just be problematic. Lethal white is a completely separate gene which does not occur in dogs. It occurs in horses, and is recessive, unlike the dominant merle in dogs. Lethal white (also known as Overo Lethal White Syndrome) kills 100% of horses which are homozygous for the gene. A lethal white will be completely white with blue eyes. It is unable to produce any pigment, like an albino, and because the cells in the intestine are linked (embryonically) to the cells which control skin pigment, they don't develop properly. The intestine is therefore contracted and lethal white foals are unable to pass faeces. This eventually results in death.

Merle Look-A-Likes

Sometimes black and white dogs with heavy ticking or roaning can be mistaken for merles. The main giveaway is that a ticked or roaned dog will have very uneven grey areas, with flecks of white showing through. Also, if the dog has tan points, they will be ticked too (if they're on the "white" areas rather than the black patches), whereas in a merle they would be solid, and the patches on a blue roan or ticked dog will be more regular than on a merle (appearing only on the back and head in the piebald or extreme white pattern, rather than all over the dog). In addition, if the dog has a butterfly nose or any blue in its eyes then it is most likely a merle as these pigment issues do not generally affect roaned dogs.
All of these dogs are merle "look-a-likes", but none of their breeds (English Cocker Spaniel, Australian Cattle Dog and Basset Bleu de Gascogne) actually carry merle:



- M: Merle (black patches on a grey base). Dilutes random sections of the coat to a lighter colour, leaving patches of full pigment. Phaeomelanin is not affected - only areas of eumelanin can be merled.
- m: Non-merle (normal expression of eumelanin).

Breeds With the Merle Allele
Australian Shepherd
Catahoula Leopard Dog
Dunker
Border Collie
Old English Sheepdog
Mudi
Cardigan Welsh Corgi
Collie (Rough and Smooth)
Shetland Sheepdog
Dachshund (known as dapple)
Beauceron
Bergamasco
Pyrenean Shepherd
Great Dane






S locus
("spotting series", affects distribution of all pigment)


White spotting on dogs is mostly determined by the genes on the S locus. When we use the term "white spotting" we simply mean white areas, not actually white spots. White spotting can occur on any colour, and will cover up both eumelanin and phaeomelanin. In technical terms this is known as epistasis. So any dog can have white markings, whether they are black, blue, liver, isabella, brindle, sable, tan-pointed, merle or whatever.

White hair occurs when the skin cells are unable to produce any pigment. The white spotting gene impairs the ability of cells on particular parts of the skin to make pigment, so the skin becomes pink and the fur white. Nails and paw pads will also become pink in areas where pigment is not produced.

Currently, only two white alleles have been proven to exist on the S locus:
S - no white
sp - piebald

A third allele may exist for "extreme white" (sw), however this has not been proven and so far all dogs with high white have been shown to be homozygous for sp instead.
The white spotting alleles are thought to be examples of incomplete dominance. This means that a heterozygous dog will express its most dominant gene, but may also be affected by the more recessive one to a lesser extent. For example an Ssp dog may have some white spotting (see below). However, the relationship between the alleles is complicated and can vary between breeds.

It has recently been shown that some dogs with white spotting do not have an sp allele at all. These are mostly dogs with "true" irish spotting (in other words, irish spotting that breeds true - this should be made clear further down the page). The allele that causes this pattern has not yet been identified and it is not known if it is also located on the S locus. For the purposes of this site we will refer to this gene as si, but remember that this may not be accurate.

Spread of White

DOG COLOR, PUPPY COLOR

Whichever white pattern a dog has, its white will always follow the same rules of spread. White starts on the farthest "edges" of the dog - the tail tip, the tip of the muzzle, the paws and the tip of the breastbone. This is known as the "trim" pattern. From there it spreads to cover the muzzle and forehead, the front of the chest, the lower legs and more of the tailtip, creating irish spotting. Next it spreads round from the front to the back of the neck, and creeps up the legs and tail. On a piebald dog, only the head, back and tail base may still be coloured. The back colouring is the next to go, followed by the tail base, then the face markings. The ears will always remain coloured unless the dog has a very high amount of white. The ears are generally the last part of the dog to turn white.

Of course the idea of white "spreading" is mainly metaphorical, to give you a picture of how white patterning works. White doesn't spread like this on one particular dog (i.e. you won't get a solid coloured puppy that gradually loses colour as it grows, until it's almost white! Although puppies do often lose or gain a little colour as they grow), it's just to show which areas remain coloured on dogs with more and more white. One way to think of it is that the dog retains colour best in the most important areas of its body - around its internal organs (body and tail base patches) and its brain (ears and face patches) - and loses colour easiest from the parts farthest from these areas. In technical terms, pigment "migrates" to different parts of the body during the development of the embryo, and the S gene determines how far the pigment migrates. Sometimes it simply doesn't reach the furthest extremities (this can be caused by a minor problem or illness during development), and this can result in a small amount of white trim on a dog without sp, for example a small chest patch on an otherwise solid-coloured dog.

The white rules aren't set in stone - sometimes individual dogs can have unusual white patterns, where, for example, the white on the legs is very uneven, or they have piebald patches in unexpected places, like on the neck or chest. However, in general, they do hold relatively true.

Residual White and White Trim



A very small amount of white on the chest, toes or tail may occur when the pigment doesn't migrate fully as the embryo develops. This is known as residual white and has no genetic basis. If a slightly larger amount of white is present then the dog may be heterozygous for sp, in other words Ssp. In a breed such as the Newfoundland you may get such a dog from crossing a "Landseer" (piebald, spsp) with a solid (SS). However, in breeds carrying piebald there is no real way to know whether minimal white markings are just residual white or indicate the presence of the piebald gene without genetic testing or test breeding, as piebald heterozygotes may have anything from a tiny chest spot to pseudo-irish markings (see below).



We can assume that the two dogs above are SS and that their markings are just residual white. This is because neither breed comes in piebald or irish spotting. If either of these dogs did have an sp or si gene then we would expect to see dogs with much more white being produced in these breeds. As it is, their white is non-genetic and breeding two dogs with white markings in these breeds will not necessarily produce puppies with any white at all.

This Staffie is a possible piebald heterozygote (i.e. carrier of the piebald allele). We cannot know for sure, but this is the most likely explanation for its white chest patch as the Staffie breed is known to commonly have the piebald gene. If this dog were bred to another sp carrier then some of the puppies may be piebalds and have much more white than either of their parents.



Irish Spotting Pattern



Irish spotting (si) is the pattern sometimes known as "boston" or "mantle", although these terms do not always refer to "true" irish spotting. On a dog with irish spotting, white is found on the legs, the tip of the tail, the chest, neck and muzzle. Many dogs with this pattern have a full white neck ring and a blaze.

True irish spotting is caused by an as yet unidentified gene, but we can assume irish spotted dogs to be homozygous for the gene (sisi) as it breeds true. This means that two irish spotted dogs bred together will produce puppies with irish spotting. We can assume that a solid dog bred to an irish spotted dog will produce a heterozygous dog with less white.

The Aussies, Border Collie and Bernese Mountain Dog shown here are all true irish spotted. None of these breeds regularly come in piebald or extreme white and their white markings breed true (implying they are homozygotes).



"Pseudo" irish spotting may look the same or very similar to true irish spotting, but is in fact not caused by sisi but by Ssp, i.e. these dogs are heterozyous piebalds. The incomplete dominance of S means that an Ssp dog may show up to roughly half the amount of white as an spsp dog. These dogs do not breed true and when two are crossed the puppies may be solid, piebald or inbetween. See below for an example of this in Boxers.



Note that not all Ssp dogs show much white, or in some cases any white at all. The amount of white on a piebald heterozygote appears to vary drastically and some may look exactly like homozygous solids. The three breeds (Staffie, Podengo Portugueso and English Pointer) all carry piebald but are not known to carry irish spotting, so these dogs are most likely pseudo-irish. A true irish spotted dog will not usually have white on the hips/knees or underside of the body, so this is another clue that sp is present.




Finally, a "flashy" irish spotted dog (one with more white than usual) may be caused by a combination of si and sp. If a true irish spotted dog also carries an sp allele then the normal white pattern may be extended. This supports the theory that si is actually on a different locus, as the two alleles appear to be inherited completely separately. This has been shown to occur in Shelties, where dogs carrying the sp allele as well as irish spotting can usually be identified by having more white around the neck and underside of the body. An spsp Sheltie has a high amount of white and is known as a "colour-headed white". Shelties are one breed known to carry both true irish spotting and the sp allele, but many breeds only have one or the other.



Piebald Pattern



Piebald (spsp) usually produces a coloured head (with or without white on the muzzle and as a blaze), and patches on the body. Generally the base of the tail is coloured, but other than that the patches may be located anywhere on the body (but rarely on the legs).




Because piebald is a recessive gene and heterozygotes piebald carriers don't always have any white markings, it can remain hidden and pop up unexpectedly. Both the Poodle (as shown here) and the Shar Pei, traditionally solid-coloured breeds, occasionally produce piebald.



Extreme White Pattern

DOG COLORS

The extreme white pattern consists of a completely or predominantly white dog with just small amounts of colour on its head and sometimes base of tail. Small body patches may sometimes be present too. Sometimes the nose is pink or partly pink, and the eyes may be blue in some breeds due to lack of pigment.
So far all extreme white dogs that have undergone genetic testing have been shown to be homozygous for the piebald gene (spsp), just like the piebalds in the section above. However, as there is a fairly large difference between those dogs and the ones shown below, it is possible there is something else going on to cause the high white. In breeds with both true irish spotting and piebald the high white may simply be caused by the interaction between homozygous irish spotting and homozygous piebald (e.g. the Sheltie). In other breeds the cause is less obvious and has led some people to postulate a further S allele - sw.

Extreme white can occasionally cause problems when it removes large amounts of pigment from the face and ears. The most common problem is deafness (due to lack of pigment in certain parts of the inner ear, which prevents it from functioning properly), but dogs with exposed unpigmented (pink) skin are also more prone to skin cancer than those with more pigment.




Split Faces and White Heads

There is thought to be a separate gene or modifier that causes some dogs with irish spotting or the piebald pattern to have a split or completely white face. A split face is when half of the face is white and the other half is coloured. This pattern occurs often on double merles, but it's just a natural part of the double merle pattern, and is not caused by any extra genes. It's only when it appears on irish spotted and piebald dogs that it raises eyebrows.



Ticking and Roan

Any white areas on a dog may be ticked or roaned due to the T gene. The ticking corresponds to the colour the area of the coat would have been if it wasn't white.

Believe it or not, the two Australian Cattle Dogs are extreme white piebalds. The solid black patches on their heads are their actual markings, and the solid appearance of the rest of the coat is created by very heavy roaning. The Large Munsterlander to the right shows heavy ticking on a piebald dog.



White Boxers

Boxers generally come in what appears to be the irish spotting pattern, so we would expect most examples of the breed to have sisi on the S locus. However, sometimes Boxer puppies are born which are completely or almost completely white. How these puppies could be regularly born to parents with much more colour perplexed Boxer breeders for a long time.

However, we can now provide an answer to this. Boxers do not have the si allele, but supposedly irish spotted Boxers are actually pseudo-irish - i.e. Ssp. When two pseudo-irish dogs are bred together some of the puppies will be homozygous piebalds (spsp).

White puppies can therefore be avoided by always breeding pseudo-irish dogs to solid dogs. A solid dog will be SS and therefore there is no possibility of heterozygous sp dogs. A solid dog will not necessarily have no white markings at all however.

The fashion for "flashy" Boxers in the show ring means that many white puppies are born. Luckily these are now usually sold as pets rather than culled.



"False" Whites

Sometimes white can occur on dogs separately to the S locus white spotting. One example is as part of the double merle pattern. A double merle will almost always have more white than its parents, and will often appear to have the piebald or extreme white pattern when in fact it does not carry those alleles. The harlequin gene also causes a similar effect. See the double merle page and merle modifiers page for more information.

White can also occur due to dilution of phaeomelanin by the I locus. Phaeomelanin is red pigment, and the I locus can dilute it to cream or sometimes white. Breeds such as the Samoyed have this second type of dilution, so they appear completely white but in fact it's not due to white spotting. They are in fact recessive red (so they cannot produce any black pigment) with dilution of their red pigment to white, resulting in a solid white dog with black nose pigment.

The main way to tell a dog with extreme white spotting apart from a dog with phaeomelanin dilution is to look at the pigment on the nose, lips and eyerims. A dog with extreme white spotting is likely to be missing some pigment in these areas, so they will be partly or completely pink. A dog with phaeomelanin dilution will have solid black in all these areas (possibly with a dudley nose, which are common on dogs with dilution).

These two dogs (a Finnish Lapphund and a Siberian Husky) are genetically black and tan (atat), but with dilution of their tan points to white. It can be easy to mistake diluted points for white markings, but points will generally be in a very regular and symmetrical pattern, with two chest spots and a vent spot, and spots above the eyes. The white will also be confined to the sides of the muzzle and not the top (except in a dog with creeping tan). The Husky has actual white spotting as well, but its cheek pattern gives it away as a black and tan.



One of these dogs is not like the others, but which is it? All are "false" whites except for one, which is an extreme white piebald. If you guessed the Staffie, you'd be right. She has pink around her eyes, ears, muzzle and underside - a sign of lack of pigment, associated with extreme whites, and a few dark spots on her ears. All the other dogs are recessive reds (ee) or clear sables with phaeomelanin dilution. Note the slight cream sheen on the coat of the German Spitz, Samoyed and Shiba, and the jet black lip and eye rim pigment on all of them. The Shiba has a dudley nose, often associated with recessive red.




- S: No white (all of coat is pigmented - no white spotting).
- sp: Piebald - homozygous causes over 50% white, with large pigmented patches on a white base). Heterozygous piebalds may have varying amounts of white, from none at all to "pseudo-irish" (phenotypically the same pattern as irish spotting but not caused by the same allele
- si: Irish spotting white on muzzle, neck, chest, feet and tail tip. This type of white spotting may not actually be on the S locus but I have included it here for lack of anywhere else to put it! Homozygous sp causes irish markings and heterozygous may have any amount of white inbetween solid and irish.

Dog Colors




T locus
"ticking series", affects distribution of all pigment)


Ticking and Roan

Ticking is flecks or spots of colour on white areas. It can occur on any white area on a dog, so long as the white is "proper" white ( i.e. so long as it's caused by the white spotting series and not by the chinchilla gene). If a dog has the ticking allele but doesn't have any white areas, there will be no visible effect.

The gene which codes for ticking has not been found yet, but it is thought to be dominant. It has been assigned its own locus - T - and there are traditionally thought be two alleles on that locus. According to the common theory, T is the dominant ticking allele, and t is the recessive clear white allele.

However, this theory does not account for the full variation in ticking and roan in dogs. It is probable that there are at least four T-locus alleles. Recent genetic research has proven ticking and roan to be distinct (but mapped to a similar area), so my own guess is that the T-series is most likely as follows:

T - ticking
Tr - roan
Td - Dalmatian spots
t - clear white

It is not clear how these genes interact or what their order of dominance is, although ticking, roan and Dalmatian spots all appear to be dominant to their absence.

Ticking amount varies greatly between dogs, and this can be partly explained by the idea of incomplete dominance. If the ticking gene (T) displays incomplete dominance over the clear white gene (t), then a TT dog would have heavy ticking and a Tt dog would have lighter ticking. It is not clear whether this is also true for roan and Dalmatian spots.

Generally, ticking is heaviest on the legs and the muzzle. If a dog has only a small amount of ticking, it will appear in these areas before appearing anywhere else. Roan, on the other hand, is more even over the whole body.

The colour of ticking/roan corresponds to the colour that the area would have been if there wasn't any white there. For example, a black-and-tan dog with white markings and ticking would have black ticking on its body and tan ticking on its legs, chest and muzzle, where it would be tan if it didn't have white. The English Setter below shows this well.

DOG COLOR

Roan is a pattern which produces heavily mottled white areas. Often only a small amount of scattered white is visible.

The three dogs are extreme examples of roan. The Australian Cattle Dog and Basset Bleu de Gascogne are genetically black-and-tan, so the roaning is black on the body and tan on the points. With the Cattle Dog, the whole dog would be white if it didn't have roaning (note there are no black patches), as it is an extreme white piebald. The overall effect on the black areas of these dogs is similar to the "salt-and-pepper" colour found on Schnauzers, except if you got a close-up look at the hairs, you'd be able to see that they're not banded like the hairs on Schnauzers are. The Basset Bleu perhaps gives a better indication that roan actually consists of lots of spots, packed very densely to cover most of the white. This has led some people to believe that roan is, in fact, just very heavy ticking. However, others believe that it is controlled by a separate gene, and recent studies appear to have confirmed this.
The German Shorthaired Pointer shows liver roan. The roan here is even more dense and difficult to distinguish from the solid liver patches on the back and head.



Black dogs with roaning often appear a greyish colour, and are commonly called "blue roans". The Cocker Spaniel above is a blue roans. Just like with "blue merles", these dogs are called "blue" but aren't actually genetically blue. The term "blue" is usually used to refer to black dogs with the dilute gene (dd), which dilutes the coat and nose to grey and the eyes to amber, but neither blue roans nor blue merles have this gene.



By contrast, these three dogs show light ticking. They are probably heterozygous for the ticking allele (Tt). As you can see, the ticking is mostly on the legs and muzzle. A dog homozygous for the ticking allele (TT) is more likely to be ticked all over.



These gundogs show heavier ticking and are mostly likely TT.



This Great Dane looks like it has ticking, but in fact the black spots are caused by the harlequin gene. It is genetically a merle, with its grey areas diluted to white. If you look carefully you can just about see a spot of actual white on its chest. This actual white is clear of ticking.

Dogs with ticking or roan are generally born white. The ticking/roaning develops as the dog grows. This can be rather dramatic, as in the Australian Cattle Dog. There is a popular myth that Australian Cattle Dogs are born white because of their Dalmatian ancestry. In fact, they are born white simply because they have the extreme white spotting pattern with roaned white areas. The roaning takes a while to develop, but the extreme white spotting is there from birth, hence the puppies are completely or almost completely white.

dog colors

Dalmatian Spots

Dalmatian spots puzzled geneticists for a long time. They are completely unique to the breed and do not occur anywhere else in the dog world. Contrary to looks, Dalmatian spots and the harlequin pattern in Great Danes are not related.

It is now fairly certain that Dalmatian spots are in fact a modified form of ticking. One of the main differences between standard ticking and Dalmatian spots is that the spots are often more sparse on a Dalmatian, bigger, and do not get more dense on the legs and muzzle. These are all probably to do with the modifier.

As well as the obvious similarity in looks between a dog with ticking and a Dalmatian, there are a few other things which indicate that spots are modified ticking:

- Dalmatians are born white and develop their spots later on.

- Dalmatians occasionally have patches on their head and/or body, as with the dog on the right above). These suggest that they have the piebald pattern, meaning that the spots can't be caused by any of the white genes. The high rate of deafness amongst Dalmatians also supports this high white piebald is sometimes associated with deafness, as in white Boxers.

- Dalmatian crossbreeds are often ticked and never, as far as I am aware, spotted. This suggests that there is a recessive modifier behind Dalmatian ticking, so the Dalmatian parent passes down one copy of the modifier, but unless it's bred to another Dalmatian, there will never be another copy of the modifier, so it cannot be expressed).

- Sometimes Dalmatians are born which display some ticking or roaning - effectively "muddying" the base white. This is probably caused by a slight mutation or error in the modifier, stopping it from being expressed normally.



- T: Ticking white areas are ticked with small flecks or spots of colour). Ticking is whichever colour would have been on that area if the dog did not have white. Suspected to be another case of incomplete dominance - a TT dog has heavier ticking than a Tt dog.
- Tr: Roan more dense than ticking
- Td: Dalmatian spots - large, round ticking spots
- t: Clear white - no ticking on white areas.




A Summary of the Summary

A locus -
Ay - sable
aw - agouti/wolf grey
at - tan points
a - recessive black

B locus -
B - non-liver
b - liver

D locus -
D - no dilution
d - dilution of eumelanin to blue or isabella

E locus -
Em - black mask
E - normal extension (no mask)
Eg - grizzle/domino
e - recessive red

G locus -
G - greying
g - no greying

H locus -
H - harlequin
h - non-harlequin

I locus -br>Alleles unknownK locus -
K - solid black
kbr - brindle
k - non-solid black

M locus -
M - merle
m - non-merle

S locus -
S - no white spotting
sp - piebald
si - irish spotting (may not be on S locus)

T locus -
T - ticking
Tr - roan
Td - Dalmatian spots
t - no ticking

A locus alleles will only be expressed when a dog does not have a dominant black (K) gene. S locus alleles (white spotting) appear on top of anything else a dog has - there is nothing that can mask them except for, possibly, I locus alleles, which may make them difficult to see. D and B locus alleles for liver and dilution will override all alleles for black (dominant or recessive) and change all eumelanin on the dog. E locus alleles are not overriden by anything except for the I (intensity) and S (white spotting) locii, so recessive red will be expressed even on a dominant black dog. Black masks will be expressed also but may not be visible on a black dog. The H locus will only be expressed on a dog with the merle allele (M locus) and the T locus will only be expressed on a dog with white spotting (S locus).


Here's a genotype for a dog which we are going to decipher to work out what the dog would actually look like:

AyatBbddeEmggHhkkmmSSTt

That looks daunting, so let's take each pair of genes individually, referring back to the summary of series above to see what each letter means:

Ayat - one allele for tan points, and one for sable. Sable is dominant over tan points, so the dog will be sable.
Bb - one allele for liver and one for normal expression of eumelanin. So the dog will not have liver pigment, as the liver allele is recessive.
dd - two allele for dilution of eumelanin. So the dog will express blue eumelanin.
eEm - one allele for recessive red, one for a eumelanin mask. Mask is dominant over recessive red, so the dog will have a mask.
gg - two alleles for no greying. So the dog will not have greying.
Hh - one allele for harlequin, one for non-harlequin. So the dog will express harlequin if it is also merle.
kk - two alleles for non-solid black. So the dog will not be solid black and will express its A-locus alleles.
mm - two alleles for non-merle. So the dog will not be merle.
SS - two alleles for no white spotting. So the dog will have no white.
Tt - one allele for ticking, one for no ticking. So the dog will have some ticking.

So our dog will have sable (which can show through because there is no dominant black allele to stop them from being expressed), it will be a dilute because it is dd, so it'll have blue eumelanin. It'll have the gene for harlequin but won't express it because it has no merle gene. Likewise, it'll have the gene for ticking but can't express that either because it has no white. It also has no no greying, no brindle, and no recessive red, but it does have a mask.

What we've ended up with is a blue sable ("fawn") with a mask:

Dog Color








DOG NOSE COLOR
DOG NOSE COLORS
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The default nose colour for dogs is Black, but a number of genes can affect nose colour.

DOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR

Liver
Liver (bb) dogs and dilute liver (isabella, dd) have noses ranging from deep brown to pink. Liver pigment does not seem to be retained in the nose as easily as black pigment, so many high-white liver dogs have completely pink noses. It is genetically impossible for a liver dog to have a black or blue nose.

Colour range in liver dogs:

DOG NOSE COLORDOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR

Colour range in isabella dogs:

DOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR

As you can see, it can be impossible to tell a liver-pigmented dog from an liver dilute just by the nose, as the colour ranges are almost identical. However, you may notice a purplish tinge to the nose of an isabella which is not usually present on a liver.

Blue
Blue (dd) dogs have noses ranging from light grey to almost black. It is genetically impossible for a blue dog to have a brown nose. Note, however, that sometimes grey dogs can occur that don't have the blue dilution gene. So an apparently "blue" dog may have a black nose and dark eyes, because in fact it is a black dog with the greying gene rather than a proper blue dilute. Sometimes blues can also be very dark, so that their coat and nose appear almost black. It can be very difficult to tell whether such dogs are blacks or blues.

Colour range in blue dogs:

DOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR
DOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR

Butterfly or Parti Nose
A "butterfly" nose is when a dog has patches of bright pink on its nose leather. These patches are randomly located and can cover any amount of the nose, from just a tiny pink spot to almost the whole nose. Butterfly noses sometimes occur on dogs with the extreme white spotting pattern (e.g. white Boxers, Bull Terriers and Dogo Argentinos), but generally they're associated with merles. Not all merles have butterfly noses though - double merles and normal merles with less dark patches than normal are more prone to them, and they are very common in harlequin Great Danes.

Butterfly noses are created when parts of the nose have no pigment (the colour pink is associated with a lack of pigment). The merle gene dilutes random parts of the pigment on the coat and nose, creating grey areas on the coat and pink areas on the nose. Harlequin Great Danes have an extra gene which dilutes the grey areas on the merle to white, so this extra strong dilution means harlequins are highly likely to have a butterfly nose.

The non-pink parts of the nose can be liver, blue or isabella if the dog is bb (liver) and/or dd (dilute). Livers and isabellas often have very light noses anyway, sometimes bright pink, so a butterfly nose may not show up on a liver or isabella merle (the whole nose may just appear pink). The first example below shows a liver merle with a visible butterfly nose.

DOG NOSE COLORDOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR

Dudley Nose
Sometimes, in specific breeds such as the Bull Terrier, the term "dudley nose" is used to describe a dog with a pink nose due to high white on the face (see butterfly nose above). However, usually it's used to describe a dog with pigment loss on its nose.
Generally the pigment loss on a dudley nose is in the middle of the nose, spreading outwards to cover almost all of the nose on some dogs. The pigment loss causes the nose to become lighter in these areas, usually ending up as a dull pink. Dudley noses never lose their pigment completely and are never as bright pink as butterfly noses or even the pink noses found on liver dogs. There is also always a darker area remaining around the edge of the nose.

Dudley noses are generally seen on dogs with solid black noses only, simply because the lighter pigment on liver, isabella and blue dogs means that areas with pigment loss are very hard to see. They are, however, very common on black-nosed dogs, including show dogs.



Snow Nose
Also known as a "winter nose", this is a dudley nose that appears during the winter months, or sometimes as a result of stress or other factors. Dudley noses are permanent, but snow noses are not.



Nose Injury
Injury to the nose can sometimes result in pigment loss, either permanent or temporary.
This dog has injured its nose, probably quite recently. You can see stripes of pink and red where the pigment has been damaged.










Dog eyes & nose color
DOG EYE COLORS
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The default eye colour for dogs is brown:
However, a number of genes can affect eye colour.

DOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR

Amber Eyes
Amber eyes usually occur when the eumelanin produced is diluted or modified by the recessive genes in the B or D series. In other words, all liver dogs (bb) have amber eyes, and so do blue and isabella dogs (dd). Occasionally dogs with black pigment also have amber eyes, but in general they are found just on livers and dilutes. Amber eyes vary from light brown (overlapping with the lighter eyes sometimes found in black-pigmented dogs) to yellow, yellow-green or grey.

DOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR

Isabella (dd) Weimaraner, showing very light eyes that almost match the shade of its coat.



Liver grizzle/agouti Siberian Husky. The pink/brown nose gives this dog away as a liver.



This English Springer shows very light amber eyes for a liver.



Blue Eyes
Genetically, there are four ways in which a dog can have blue eyes. Three of these are linked with pigment loss in the coat.

DOG NOSE COLORDOG NOSE COLORDOG NOSE COLOR

The most common way is as a side effect of the merle gene. Merle dilutes random parts of the pigment, including the eyes and nose. This sort of dilution causes blue colour in the iris (contrary to common belief, animals with no pigment around their eyes do not always have pink or red eyes like albino rodents do - lack of pigment or very diluted pigment often results in blue eyes, as it does in albino humans). Because of the random pigment loss, often merle dogs have "butterfly" noses (see nose page) and blue, wall or split eyes. Wall eyes are when a dog has one blue eye and one brown or amber eye, and a split eye has some blue in it and the rest is brown or amber. Split eyes vary from mostly blue to mostly brown or amber.

The more dilution there is in the coat of a merle (i.e. the more grey/diluted areas), the more likely they are to have blue eyes or a butterfly nose. A heavily merled dog (large areas of black or liver) is unlikely to have either of these traits. Double (homozygous) merles are highly likely to have blue eyes and a completely or almost completely pink nose because of the combination of merle dilution and large amounts of white around the face.
This red (liver) merle Aussie has one amber eye (because of its bb liver pigment) and one blue. If you look carefully you can see a sliver of blue in the amber eye.

DOG NOSE COLOR

The merle Cardigan Welsh Corgi shown here has two solid blue eyes.

DOG NOSE COLOR

This merle Border Collie has very striking light blue eyes, which is surprising as its merling is fairly heavy.

DOG NOSE COLOR

The second way in which blue eyes can occur is when a dog has large amounts of white around its eyes. White areas on the coat are where the cells are unable to produce any pigment, so if these areas spread to the face then there may be pigment loss in the eyes and on the nose, making the nose pink and the eyes blue. This only tends to occur on very high-white dogs with the extreme spotting pattern, such as white Boxers, and even then is fairly unusual.

The third way is when a dog is affected by the C series. The C series is albino. There are no confirmed cases of true albinism in dogs, however "white" Dobermanns have a very light coat (with the main coat appearing as a very light isabella colour and the tan points light cream), blue eyes and a fully pink nose, and this is thought to be a form of albinism.

Lastly, blue eyes can be inherited as a completely separate gene, unaffected by coat colour. This gene is, however, rare. It is rumoured to occur occasionally in the Border Collie, but mainly it's seen in the Siberian Husky. Huskies can have one or both blue eyes, regardless of their main coat colour, ranging in shade from almost white to sky blue. This is particularly striking when seen on black dogs.

These two Huskies have almost identical coat colours, but one has two deep brown eyes and the other has one blue eye (eyes of different colours are known as "wall eyes"): As you can see, both dogs have full black nose pigment and black around the eyes - eye rims, so the only explanation for the blue eye on the dog on the right is that it is inherited separately to coat colour and pigmentation.

Dog Eyes Colordog eye color

A Husky with two blue eyes.



This mixed breed has wall eyes, but is not a merle (the greyish areas are roaned white). This means it must have the independent blue eye gene, which is most likely from a Border Collie or Husky parent. The large amount of Husky crosses with blue or wall eyes suggests that the independent blue eye gene is dominant, in Huskies at least.

DOG COLOR NOSE

The recessive blue eye allele also appears to crop up in Beagles such as this one. Note the jet black eyerim pigment and the colour surrounding the eyes, proving that the blue eye is not due to white spotting.

This photo is (c) by Darlene of AlaDar Beagles








Dog coat \ fur color
DOG COAT COLOR TERMS
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DOG COAT FUR COLOR

DOG COAT FUR COLOR


The frustrating thing about dog genetics is the vast array of different names given to the same colours in different breeds. For example, genetically recessive red Labradors are "yellow" but Golden Retrievers are "golden". You can't have a golden Labrador or a yellow Golden Retriever, yet they are genetically the same colour! This can be a bit of a barrier when it comes to working out the genetics of particular breeds, so to make things easier, here's a list of some of the terms you will find (either on breed standards or just being used by breeders), and what they actually mean in terms of the genetics we've studied on this site.

Just a diagram with some of the main terms used to describe body parts in dogs. This may help if you're trying to interpret breed standards.



Term

Breeds

What it means

Albino

Dobermann

Not thought to be real albino, just a dog with very, very diluted pigment

ASCOB

American Cocker Spaniel

Any solid colour other than black

Ash

Various - general term

Isabella (liver dilute, bbdd) or very washed-out red

Banding

Various - general term

When hairs have bands of more than one colour. Associated with agouti (aw)

Beauty spot

Various, but mainly Cavalier King Charles Spaniel and King Charles Spaniel (English Toy Spaniel)

Coloured spot in the middle of the forehead blaze on a piebald dog

Belge

Affenpinscher

Sable with black tipping

Belton

English Setter

Roan or heavy ticking in any colour

Bi-black

Shetland Sheepdog

Black and white with no tan markings

Bi-blue

Shetland Sheepdog

Blue merle and white with no tan markings

Biscuit

Samoyed, American Eskimo

Cream (appearing either as patches or on the whole of the dog)

Black and silver

Various - general term

Black with tan points (atat) which have been diluted to off-white by the Intensity gene

Black brindle

Various - general term

Heavily brindled dog (i.e. with thick black stripes, so very little red shows through)

Blenheim

Cavalier King Charles Spaniel and King Charles Spaniel (English Toy Spaniel)

Deep, clear red (recessive red, ee) with white markings in the piebald pattern. The same colour as the Red and White Setter

Blue fawn

Basenji, Rat Terrier

Sable with blue (diluted black, dd) pigment, or sable with isabella (diluted liver, bbdd) pigment, giving a grey/blue cast to the coat

Blue roan

Various - general term

Black-pigmented dog with roaned white, which appears a bluish colour

Boston

Great Dane, Boston Terrier

Black (or brindle) dog with white markings in the irish spotting pattern

Brindlequin

Great Dane

Harlequin with brindle patches

Bronzing

Various - general term

A brownish cast to the coat of a longhaired black dog, usually caused by sun bleaching

Butterfly nose

Various - general term

Partly pink nose on a merle

Charcoal

Various - general term

Very dark blue dilute (dd), often appearing almost black

Check

Boxer

Brindle or fawn (sable) boxer with the piebald white pattern

Chocolate

Various, but particularly the Labrador

Liver (bb)

Clear-coloured

Various - general term

One solid colour all over (i.e. no sabling on a red dog). Clear-coloured dogs may still have white markings (white is not a colour)

Copper

Siberian Husky

Reddish liver (bb)

Creeping tan

Various - general term

Pattern inbetween traditional tan points and saddle. Generally the tan covers the whole of the muzzle and eyebrows, most of the legs, and the front of the chest. Probably caused by a modifier that extends the normal tan points

Cryptic merle

Various - general term

Merle dog with little or no visible merling, i.e. one which appears solid black (or liver, etc)

Dapple

Dachshund

Merle

Dark merle

Various - general term

A merle dog with a dark grey base coat (rather than the usual light grey). This is caused by intermingled black hairs

Deadgrass

Chesapeake Bay Retriever

Light liver-pigmented red. A duller shade than sedge.

Dilute spot

Various - general term

Patch on a merle which is lighter than the other dark patches

Domino

Afghan Hound

Usually a dog with a greyish/brown back and cream/white underparts, caused by the grizzle gene (Eg)

Dorsal stripe

Various - general term

Dark line along the back, often caused by sabling

Dudley nose

Various - general term

Pink or liver nose

Fawn

Various - general term

Clear sable, usually with a black mask

Fawn

Dobermann

Isabella (dilute liver) with tan points

Fawnequin

Great Dane

Harlequin Great Dane with sable (fawn) patches rather than black. Black patches may still appear on the muzzle, where most Danes have black masks

Fawn merle

Various - general term

Sable merle. In clear sables, merle may only be visible on the mask (if present)

Flashy

Various - general term

A dog with more white than normal for its breed, or a dog with clear white (no ticking)

Flecking

Various - general term

Ticking

Flowered

Shar Pei

Piebald

b>Ginger merle

Cardigan Welsh Corgi

Brindle merle with an orangey base coat

Grizzle

Saluki

Black/brown with tan points/underparts. Caused by grizzle gene Eg, as in domino Afghan Hounds

Grizzle

Terriers

Intermingled brown and black hairs (usually on a saddle)

Harlequin

Australian Shepherd

Merle dog with white patches in its merling (caused by an unidentified modifier - probably the same as tweed)

Harlequin

Beauceron

Blue merle with tan points

Harlequin

Great Dane

Merle modifier which dilutes the grey parts on a merle to white, leaving jet black patches on a white base. Overall appearance is similar to a Holstein cow

Havana Brown

Havanese

Black brindle (brindle with very thick black stripes). When the coat grows in, such dogs can appear to be a brownish colour, like liver but without liver pigment

Isabella

Various - general term

Dilute liver (bb on the B locus and dd on the D locus)

Isabella

Shar Pei

A light, dullish red dog with a blue mask and blue sabling (so a blue dilute sable, dd)

Isabella

Siberian Husky

Off-white (light cream), caused by dilution of phaeomelanin (red). May have a black or liver saddle.

Ivory

Various - general term

A solid white dog with a slight cream sheen

King Charles

Cavalier King Charles Spaniel and King Charles Spaniel (English Toy Spaniel)

Black with traditional tan markings (atat)

Lavender

Shar Pei

Light purplish tongue colour on a dilute Shar Pei

Light red (and red)

Siberian Husky

Light reddish liver (bb). Dark liver is called copper in this breed.

Lilac

Shar Pei

Isabella

Lion

Ibizan Hound

Light red

Maltese blue

Various - general term

Blue (diluted black, dd)

Mantle

Great Dane

White markings in the irish spotting pattern (muzzle, neck, chest, lower legs and tail tip)

Mask

Various - general term

Black (or liver, isabella, or blue) on the muzzle area (may include the eyebrows and the ears), caused by the Em gene

Merlequin (aka merlekin, merlikin)

Great Dane

Double merle. The double merle causes broken patches of merle on a white base, giving a pattern similar to harlequin (but more regular)

Minimal merle

Various - general term

Merle dog with very little visible merling. In effect, an almost completely black (or liver, etc) dog with just a small patch of diluted fur

Mosaic

Various - general term

Generally a red dog with random black sections on its coat, caused by a very rare somatic mutation

Mosaic spot

Dalmatian

A red or tan spot on an otherwise black-spotted Dalmatian. Caused by a mutation

Muddy merle

Various - general term

Blue merle with a brownish sheen to the grey parts of the coat

Mustard

Dandie Dinmont Terrier

Light tan (probably sable) with white furnishings and head

Onyx brindle

Various - general term

Black brindle (thick black stripes)

Particolour

Various - general term

Any dog with two distinct colours in its coat

Pearl

Rat Terrier

Isabella (diluted liver, bbdd)

Pencilling

Various - general term

Black lines on the toes of a tan-pointed dog

Pepper

Dandie Dinmont Terrier

Black with tan points (atat) diluted to white by the Intensity gene. The black is diluted to silver by the progressive greying gene (G). Appearance is of a light grey dog with white underparts.

Phantom merle

Various - general term

Dog which has the merle gene, but is solid red because of the recessive red gene (ee) or clear sable. As there is no visible merling, it can be impossible to tell the dog is a merle

Pinto

Akita

Piebald

Pips

Various - general term

Tan markings above the eyebrows and on the cheeks on a tan-pointed dog

Platinum

Various - general term

Silvery white

Points

Various - general term

Markings (usually tan, or a variant of) on the sides of the muzzle, the neck, chest, eyebrows, lower legs and vent

Porcelaine

Great Dane

Harlequin with patches of any colour other than black

Powder merle

Various - general term

Pale-coloured merle with only a few dark patches

Prince Charles

Cavalier King Charles Spaniel and King Charles Spaniel (English Toy Spaniel)

Tricolour (black with tan points and white markings)

Red speckle

Australian Cattle Dog

Red roan

Reverse brindle

Various - general term

Black brindle (very heavy black striping)

Ruby

Cavalier King Charles Spaniel and King Charles Spaniel (English Toy Spaniel)

Deep, clear red (recessive red, ee)

Sable

German Shepherd Dog

Agouti or wolf grey (aw)

Salt and pepper

Schnauzer

Wolf grey (agouti, aw, with very strong dilution of phaeomelanin, turning the red to white). Appears as a grey dog with white points in the traditional tan point pattern. On longer fur, banded hairs are visible

Seal

Various - general term

Appears black with a brownish tinge. Genetic basis is unknown.

Sedge

Chesapeake Bay Retriever

Liver-pigmented sable, sometimes with a liver mask

Sesame

Shiba Inu

Shaded red sable with urajiro markings

Silver brindle

Various - general term

Brindle with greyish stripes (but not a blue brindle - silver brindles are not dilutes)

Slate

Various - general term

Dark blue (which is diluted black, dd)

Slate merle

Various - general term

Merle with diluted pigment, dd (i.e. a proper 'blue' merle!)

Sooty

Various - general term

Sable with black sabling "muddying" the red. Sometimes called a sabled fawn.

Spectacles

Keeshond

Cream colour around the eyes on a wolf grey dog

Splash coat

Siberian Husky

Piebald

Tiger brindle

Various - general term

Light brindle with sparse stripes

Tricolour (tri)

Various - general term

Literally "three colours". Usually a black and tan-pointed dog with white markings (to any extent), although liver, blue and isabella tris occur too

Tricolour

Terriers and hounds

Tan dog with a black saddle and white markings (to any extent)

Trim

Various - general term

A small amount of white on the chest, muzzle, toes and/or tail tip

Trindle

Various - general term

Brindle tricolour (i.e. black with brindle points and white markings. Brindle never appears as the main colour on a tan-pointed dog)

Tweed (tweed merle)

Various - general term

Modified merle, causing patches of brown and tan to appear on a blue merle.

b>Urajiro

Shiba Inu and others

White markings on the underside, roughly following the same pattern as tan points. Urajiro is caused by a specific gene and affects only phaeomelanin (red)

Vent marking

Various - general term

Tan patch under the tail on a tan-pointed dog

White collar

Various - general term

White reaching all the way around the neck

Widow's peak

Various - general term

Brown or black marking running down the middle of the forehead on a shaded sable dog

Wolf grey

Various - general term

Agouti (aw) with phaeomelanin dilution turning the base coat to grey and the points to cream, leaving some black on the back. This is the colour of the Keeshond and Norwegian Elkhound.

Apricot
Buff
Chestnut
Cinnamon
Cream
Deer
Fawn
Fallow
Gold
Lemon
Mahogany
Orange
Red
Russet
Rust
Sandy
Stag
Tan
Tawny
Wheaten
Yellow

Various

Used to describe various shades of red, from the lightest (cream) to the darkest (mahogany)










WEIRD STRANGE ODD DOG COLORS
DOG COLOR ODDITIES
(Strange Colors)




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Somatic Mutations and Chimera

A somatic mutation is a mutation that occurs in the body cells after the embryo has begun to form. Cells are divided into two categories - germline cells (i.e. gametes) and somatic cells (all other body cells). When a mutation occurs at the point of conception (when the first cell begins to replicate) then this mutation may enter the germline and so be passed on to future generations, but after this point the two types of cell replicate separately and any mutations occuring in the somatic cells will only be passed on to the descendents of those cells and will not be passed to the next generation. This is because only the information carried in the gametes will go on to form a new individual - of course, mutations may well happen in the gametes too during the individual's lifetime, and these mutations will be passed on.

This may sound complicated, so perhaps an example will make it clearer. Cancer is a somatic mutation. Sometimes a mutation occurs in a cell that makes the cell "immortal". Most cells can only replicate a certain amount of times, but when this particular mutation happens the cell is able to replicate itself indefinitely - this is the biological definition of immortal, and this is how tumours form and also why they can be very difficult to get rid of. However, although the predisposition to cancer may be genetic, cancer itself is not as it occurs as a mutation in cells which will not go on to form new individuals.

Another type of somatic mutation in dogs involves pigment. If a pigment mutation takes place during the development of the embryo, patches of another colour may appear on the puppy. This is most often seen as black patches on recessive red Labradors and Golden Retrievers (it is not certain why it seems to appear in these breeds more than others). The painting at the top of the page shows what this often looks like. All it takes is for one cell to mutate early on in development - then all cells descended from that one as the embryo grows will be the different colour. Due to the mutation these patches effectively have a different genotype to the rest of the dog, and this is sometimes known as mosaicism.


DOG COLOR

A very rare form of mosaicism is the chimera. A chimera is a single animal that is the combination of two separate embryos. You may have heard this term used to describe creatures with the DNA of two different species spliced together, such as human/mouse embryos created in labs, but it also occurs naturally when two fertilised eggs combine very early in development. Effectively, the resulting animal is two individuals in one. It has patches of one set of DNA and patches of the other as well. Sometimes it can be impossible to tell a chimera without genetic testing (particularly in humans), but in some cases the resulting animal may be a combination of two different colours. I do not know of any proven dog chimeras, but the effect may be similar to the somatic mutations already discussed, but more extensive (roughly 50/50 of each colour in random patches).

This incredible Labrador appears to show somatic mutations, but is a possible chimera due to the extent of his patching. These photos were taken at a rescue centre in France and sent to me by Damien Ricco but we have been unable to locate the original photographer.




Vitiligo

Vitiligo is a condition where the skin/coat cells stop being able to produce pigment, causing areas of white. It can have a variety of causes (including an auto immune disease that attacks pigment cells), but in dogs does not usually have any adverse effects other than coat colour. The pigment loss is generally concentrated around the head/face, but often spreads to the rest of the body.

Phaergus, the Newfoundland above, has developed what appears to be vitiligo on his head, chest, back and tail. The first picture shows Phaergus in May 2010 (at 3 years old), and the second shows him in March 2011. Phaergus and his photos belong to Bobbi Walker.




Greyhound Spots

I have not seen this phenomenon in any other breeds but it appears to occur fairly frequently in black Greyhounds. There are small white spots on the coat, almost like ticking in reverse. These dogs look like they have been sprinkled with snow. It is not clear whether this is caused by some form of greying, vitiligo, or whether it is actually a colour gene. Vitiligo is unlikely as this is usually concentrated around the face and not in small spots all over the body.




Other Strange Things

This unusual pattern appears to occur only in Finnish Lapphunds. Some are darker than this - but all have the tan spectacles around the eyes. It does not fit any of the usual tan patterns, although the spectacles are reminiscent of agouti.


STRANGE WEIRD UNUSUAL DOG COLORS

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Extreme colored dogs worldwide (photos)

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

DOG COAT, FUR, WOOL, HAIR

STRANGE DOG and PUPPY COLOR
STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR
STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR
STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR
STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR

STRANGE DOG and PUPPY COLOR













STRANGE DOG and PUPPY COLOR








DOG EYE AND NOSE UNUSUAL STRANGE COLORS
HEALTH PROBLEMS
RELATED TO COLOR

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There are a few colour genes which can occasionally cause health problems in dogs, most notably merle.

Double Merles and High Whites
A homozygous (or "double") merle is one with two copies of the merle gene, and this severely impairs its ability to make pigment, leaving large areas of the dog pigmentless (white). Pigment is actually necessary for certain parts of the body to function correctly, so lack of pigment can cause health problems. Dogs with large amounts of white caused by the homozygous piebald allele (sp), such as Bull Terriers, Boxers and Dalmatians, can also have some of the same health problems as double merles, particularly deafness (which is a huge problem in Dalmatians).

Lack of pigment in particular parts of the inner ear can cause deafness, which can be unilateral (just one ear) or bilateral (both ears). It is commonly claimed that dogs with white ears are always deaf, but in fact it's been shown that whether or not pigment is visible on the outer ear does not affect whether or not the dog can hear. In other words, a dog may have coloured ears but still be deaf, and a dog with white ears will not necessarily have any problems.

The double merle gene can also cause eye deformities. This is because the location of the eye cells in an embryo happens to be the same place that pigment starts to appear. If there is a problem with the pigment, this can therefore affect the development of the eyes. Problems include irregularly-shaped pupils, subluxated pupils (not positioned in the right place), microphthalmia (abnormally small eyes, usually with impaired vision), and other, less visible abnormalities causing blindness and bad vision.

Lack of pigment anywhere on the dog can make the skin much more sensitive to the sun. This is a particular problem on the nose, as it is so exposed, but any area of pink skin is susceptible to sunburn and skin cancer. The same problem occurs with any animal which has little or no pigment. White cats are probably the most well-known example. Skin cancer rates in white cats are extremely high and a surprising proportion of cats with white ears end up having their eartips amputated to stop the spread of cancer. The main way to prevent sunburn in animals is the same as with humans - apply suncream!


Dilutes
There is a common misconception that dilutes are in some way naturally sickly - this is not in fact the case. The dilution gene does impair the ability of the cells to make pigment, but only in that it causes the pigment that is made to be less intense. As with most recessives, the dilute allele is in some way "faulty", but it is only faulty in its ability to produce full-strength eumelanin.
The ability or inability of the cells to produce full-strength eumelanin does not affect the health of the dog, simply its colour.

That said, the idea of dilutes as unhealthy most likely has its foundations in Colour Dilution Alopecia. This is an apparently genetic disease causing hair loss and skin problems. A dog with this disorder will typically appear "mangy" and have partial hair loss. It is usually reported from blue dogs, particularly Dobermanns, but presumably it affects isabella dogs too diluted livers. Any colour can carry CDA or be homozygous for it, but only blues and isabellas will have symptoms.

DOG EYE AND NOSE UNUSUAL STRANGE COLORS

CDA does not occur on all dilutes and its frequency varies between breeds. It is particularly common in Dobermanns, occuring in up to 80% of dilute dogs. Dilutes in other species such as mice are caused by the same gene, and yet CDA is not known in these, implying it is not an unavoidable consequence of dilution. It is thought that CDA may be caused by a specific dilution gene - labelled dl. Just as there are various different b alleles that all cause the liver colour (phenotypically the same, so only distinguishable through genetic testing), it is probable that there are a number of different d alleles as well, and only one of these causes CDA.
What this means is that CDA is most likely caused by a recessive allele but could theoretically be bred out of most lines by careful breeding and genetic testing.

This blue German Pinscher appears to have mild alopecia. Its coat is dull rather than having a healthy shine, and it seems thin and patchy.

DOG EYE AND NOSE UNUSUAL STRANGE COLORS

The same problem can also occur (albeit rarely) on black or liver dogs, and is known as Black Hair Follicular Dysplasia. It affects black/liver hairs only, leaving all other hairs as normal. Because this condition is so rare, it often goes undiagnosed. I used to know a Jack Russell Terrier mix who was white except for a black patch on his back, which was hairless. His condition puzzled a whole string of vets and skin specialists, who suggested various types of mange and allergies, and he was never properly diagnosed as having Black Hair Follicular Dysplasia. Unfortunately for dogs with genetic hair loss conditions, there is no cure, although these conditions do not generally cause the dog to be itchy or uncomfortable and so are mostly harmless.








DOG EYE AND NOSE UNUSUAL STRANGE COLORS
BASIC DOG COAT COLOR GENETICS
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DOG FUR COLORDOG COAT COLOR
ADVANCED DOG COAT
COLOR GENETICS

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Two Different Types of Pigment
The key to understanding dog genetics is simply this: there are two types of pigment which create coat colour in dogs (and most other mammals). Pigment is just the thing that gives each strand of hair its colour, just like pigment in paint or dye, or pigment in your own hair. All coat colours and patterns in dogs are created by these two pigments, which are both forms of melanin. Each of the pigments has a "default" colour, and it can then be changed by various genes.

1. Eumelanin



Eumelanin is, by default, black pigment. All black areas on a dog are caused by cells producing eumelanin. However, there are genes which turn eumelanin into other colours - liver (brown), blue (grey), or isabella (a dusty pale brown). If a dog has any of the genes to turn its black eumelanin into liver, blue or isabella then all of the black in its coat will be changed. This is because these genes restrict and/or alter the production of eumelanin, so the cells aren't able to produce full-strength pigment. We call blue and isabella dogs "dilutes" for this reason. They either lack the genes which tell their cells how to produce proper eumelanin, or their cells know how to produce it but can't.

As well as being found in the coat, eumelanin is present in the other parts of the dog that need colour - most notably the eyes (irises) and nose. The nose will be black, liver, blue or isabella depending on the type of eumelanin the dog can produce. The colour in irises is produced by layers of pigment, and brown eyes like most dogs have (and like many people have too) are caused by black eumelanin in those layers. When a dog has altered/restricted production of eumelanin, the irises are also unable to produce full-strength eumelanin. This means that the dark pigment in the eye becomes lighter, and the eyes turn into a light brown colour, known as amber or gold. This isn't really a colour we find in human irises, so it's hard to illustrate it with an example. The closest we have are hazel eyes, which are a mixture of green and brown, and so appear lighter than normal brown eyes.

DOG COLOR

This shows the variation in colour of eumelanin. Black is the default, then it can be turned into liver by one set of genes, and black and liver can be turned into blue or isabella by another set of genes. Blue is diluted black and isabella is diluted liver.

When we talk of dogs that are "black pigmented", "liver pigmented", etc, we mean that is the colour of eumelanin that the dog can produce. Sometimes these dogs have no eumelanin at all in their coats (their skin cells produce only the other type of pigment, phaeomelanin), but we can tell what their "pigment colour" is by looking at their nose. A black nose means the dog produces black eumelanin, and so on. It's confusing to talk of a dog's "pigment colour" like this, because as we know, eumelanin isn't the only type of pigment. But it's common practice, and I'll be using such terms a bit on this site.


2. Phaeomelanin



The second type of pigment, in some ways less important than eumelanin, is phaeomelanin. This is red pigment. The term "red" covers everything from deep red (like Irish Setters) to light cream, encompassing gold, yellow and orange. Whenever we talk of red, unless we're talking particularly about Setters, we mean the whole range of tan colours.

Phaeomelanin is produced only in the coat. It does not occur in the eyes or the nose, so any genes which affect the colour/intensity of phaeomelanin will not affect the eyes or nose. Only eumelanin occurs in those areas, and so only genes which affect eumelanin can affect the eye or nose colour.

This shows the variation in colour of phaeomelanin. Unlike eumelanin, it doesn't occur in two distinct colours (black and liver, with dilutes counting as shades of those), but rather just one colour, which varies in intensity. The most intense phaeomelanin colour is Irish Setter red. The default colour is probably golden, with different genes causing it to be more or less intense (i.e. telling the cells to produce a higher density of pigment particles, so making the colour stronger, or a lower density, making the colour weaker, so lighter).
So far so good, but this doesn't seem to explain all the coat colours in dogs - how about white?


White

White isn't really a colour, so white hair on animals isn't caused by pigment but a lack of pigment. In dogs it is a lack of both eumelanin and phaeomelanin. White areas on animals are simply caused when the cells cannot or do not produce any pigment at all. Sometimes the whole animal is affected, like in albinos, and sometimes just parts of it are affected, like in dogs with white markings. It can affect the production of eumelanin in eyes and noses too, turning noses pink and eyes blue (or red in proper albinos). There is also a second type of white, which is caused by dilution of red (phaeomelanin) pigment, making the cells produce fewer pigment particles than normal, so the colour gets lighter. If it is diluted enough, it can become white. Many white dogs have a slight ivory/cream sheen to their coats because their cells are still producing a very small amount of pigment. This sort of white does not usually affect eumelanin, so any black/liver/blue/isabella areas on the coat will stay dark, and the eyes and nose will do too.


DOG COLOR

Distribution of Pigment

The colour genes in dogs do two things - they determine the eumelanin and phaeomelanin colours/shades, and they control the distribution of these two pigments. They tell certain cells to produce eumelanin, others to produce phaeomelanin, and sometimes they tell them to not produce pigment at all. Exactly which cells are told to produce what is determined by the exact set of genes, although it can be random to a certain degree (e.g. puppies may have slightly different white markings to their parents, or patches in different places). Sometimes genes can even tell cells to switch which type of pigment they are producing every once in a while. This means that as a hair grows, it becomes banded with black and red, because the cell produces black (eumelanin) for a while, then changes to red (phaeomelanin), then back to black, etc. It's a bit like when you highlight your hair and after a while the roots start to show through. The overall colour of an animal with this sort of black and red banding will generally be a muddy brown from a distance, and close up you will be able to see the black parts of the hairs. It's called agouti, and it's the colour of wild rabbits and mice, as well as a large amount of other mammals. It's popular amongst wild animals because it provides very good camouflage. It also occurs in dogs, but it looks a bit different, like the colour of a wild wolf rather than a rabbit and isn't very common.








DOG BREEDING IN COLORS
BREEDING IN COLORS
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Getting Started
As we've discussed already, every dog has a set of locii (points on its strand of DNA) with two genes on each one. For example, on the B locus they might have the genes B and b, b and b, or B and B. These are picked from a list of possible genes for each locus, which for the B locus is just simply B and b, but for other locii there may be a longer list (the A series, for example, has five possible genes - Ay, aw, as, at and a).

When dogs breed (and every other mammal, including humans), each parent passes on just one gene from each of its locii. This gene is picked at random, so each of the two genes has an equal chance (50%) of being passed on. However, it isn't just a case of the same gene being passed on to all the puppies.

When a sperm cell fertilises an egg, an embryo starts to develop. The sperm cell is just one of thousands that have been produced by the male dog. Each sperm cell is an individual. It carries half the information needed to create a new life, and that information consists of one set of genes. This means it has one gene from each of the male dog's locii, and which of the two genes that is is entirely random. There are lots of different genes, so the exact combination carried by the sperm cell may well be pretty unique.

Egg cells are exactly the same - they also carry one set of genes from the parent - one from each pair. However, fewer of them are produced, as, obviously, it wouldn't be good to have more than a certain amount fertilised at the same time.

Just a few out of the thousands of sperm produced will actually reach the eggs in the female dog's womb. The rest will die. The ones that do reach an egg will join with it and insert their nucleus, which is the part of the cell that carries the genetic information (in this case, the incomplete strand of DNA), into the egg. The nucleii of the egg and the sperm then join together to create a new nucleus which carries all the necessary genetic information to develop into an embryo.

So we can see that genetics involves a lot of random chances. Because of this, it's hard to be certain of anything when you breed a pair of dogs. We can't say that the puppies will definitely be this colour or that pattern. All we can do is predict the chance that the puppy will be a certain colour. It's important to remember this when reading the rest of this page. When predicting litters we usually give at least four possible outcomes for the puppies. This does not mean that you will get four puppies and one will be each of those colours. All it means is that each puppy has a 25% chance of being each colour. Occasionally you may even end up with each puppy in the litter inheriting the same genes and so being the same colour; it just depends on which sperm reach the eggs. It's like throwing dice. Sometimes you may be unlucky or lucky, if you're playing a board game! and get six sixes in a row. That seems really unlikely, but actually, each time you throw the die, the chance of getting a six is the same. Each throw is independent of all the others (the chance of getting a six the second time is not dependent on whether you got a six the first time), and genetics is very similar. Just because only one in six of the puppies are supposed to pure white according to our calculations, it doesn't mean that if the first pup born is white, the second is any less likely to be white too!


Labrador Case Study



Now that we have a basic idea of how breeding works, let's look at a case study.
Let's say we have lots of Labradors, and we are going to breed them. Not in some bizarre, frenzied dog orgy, of course, but a pair at a time.
First, we decide we want some black puppies, so we pick two lovely, shiny black dogs. Let's call them Blackie and Sooty, because we're uninventive.



We breed Blackie and Sooty, and we get this litter:



Well that all seems to be in order. We bred a black dog with a black dog and we got black puppies. Seems to make sense, doesn't it?

Enthused by the success of our first litter, and anxious to breed some more black puppies to sell to the black Lab-loving public, we choose two more black dogs: Jet and Ed.



We breed Jet and Ed and get this litter:



Looks good... but oh! Hang on a sec! One of those puppies is brown. How on earth did that happen?

Our surprise at getting this chocolate-coloured puppy leads us to look up dog colour genetics on the internet. It turns out that whether a dog is black or liver (chocolate/brown) depends on the genes it has on a particular locus - the B locus. There are two possible genes - B is black, and it's dominant over b, which is liver.
Here's a list of the possible genotypes and what dogs with those genotypes would look like:
BB - two copies of black, so puppy will be black.
Bb or bB - one copy of black, one of liver. Black is dominant, so puppy will be black.
bb - two copies of liver. Puppy doesn't carry any black, so it must be liver.


Punnet Squares
The mystery of the chocolate puppy can be solved by working out the genotypes of the parents. We can see from the list above that if a dog is black, it must be BB or Bb. If we calculate some breeding results from all the possible crossings of BB/Bb dogs, we can work out which breeding/s will produce chocolate puppies, and so from that we can tell the genotypes of the parents.

The quickest and easiest way to calculate breeding results is to use a Punnet square. Firstly we'll see what happens when we breed a Bb dog with a BB dog.


This is a blank Punnet square, ready for us to fill in. As you can see, there are two rows and two columns. The number of cells in the table, not including the ones we will put the column and row titles i is four, so we'll get four puppy options each time we fill it in.


First we put the genotype of each parent in the boxes at the top and the side (the title cells). We put just one gene letter in each cell, and it doesn't matter which parent goes across the top and which down the side - it makes no difference to the results.


Now we need to fill it in. Each of the four cells represents one possible puppy genotype. To work them out, we trace up from the cell to the column title and across to the row title. It doesn't matter which order we do this in, whether we look at the row or the column first, but I usually do column then row.



Here's the finished Punnet square. If we refer back to the list above, we can see that all the puppies in this litter would be black. There are no bb puppies, and they're the only dogs that can be liver. So we haven't found Jet and Ed's genotypes yet. We need to keep looking!

Crossing a BB dog with another BB dog will obviously only produce BB puppies, so Ed and Jet cannot both be BB. From the example above we now also know that it can't be right that one of them is BB and the other Bb. So the only other option we can try out is Bb x Bb:


So that's three black puppies (BB, Bb and bB), and one liver! It looks like we've found what we were looking for. Because the liver gene (b) is recessive, it can be hidden, which means a dog can carry one copy of it without expressing (displaying) it. That means that two dogs carrying liver can be black themselves, but produce liver puppies. It's a bit of a problem when it comes to breeding, actually, because recessive traits, such as liver and dilution, can remain hidden in lines for many generations, then suddenly crop up when a dog carrying the trait is bred to another with it (if the gene is very rare in the breed then it can be a long time until this happens, if it ever does). This is why breedings sometimes throw complete surprises, like silver (blue) Labrador puppies in a breed which, to all intents and purposes, contains no silver at all. That one lone recessive silver gene (d, on the D locus) has been passed down from generation to generation, completely unknown to the breeders, until finally it's met another one. It might have come from a cross breeding with another breed many years ago, which doesn't show up on the pedigrees and no longer has any effect on the look of the dog, so all the dogs in the line look exactly like normal Labradors. Such rare recessive traits can be impossible to eradicate from a breed, simply because you can't tell which dogs carry them. In recent years, however, genetic testing has helped to identify the carriers.

Anyway, so now we know that Jet and Ed both have the genotype Bb. However, if we wanted to get liver puppies in the future by just breeding black dogs, how would we know which black dogs are Bb and which are BB, without doing any genetic testing? Luckily for us liver is quite common in Labs, so it's not like trying to find out which Labs carry silver, where you could do hundreds of test breedings and still not get silver pups because the chances of finding another dog with the gene are so slim. The way to tell if a black dog is Bb or BB is to breed it to a liver.

We can be certain that the liver parent will be bb, because it can't be anything else. If the black parent is BB, we'll end up with all black pups, because each pup can only possibly inherit B from the black parent. All of the puppies will have the genotype Bb. But if the black parent is Bb, half of the puppies will be liver. How do we know this?

By doing a quick Punnet square:

Dog colors

The resulting puppies will be:

Dog colorsDog colorsDog colorsDog colors
So if any of the puppies in the litter are liver, the black parent must be a carrier of of the liver gene, so it must have the genotype Bb.

Lastly, it's important to note that due to the nature of recessive genes, the only breeding which will produce all liver puppies is bb x bb, so two liver parents.

Dog colorsDog colorsDog colors Dog colors








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