So long! Helka’s litter
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Home » Appearance of the pumi

Pumi colours

Submitted by on Nov 30, -0001 – 12:00 AMNo Comment

What is colour?

Colour has been an integral trait in the development of many dog breeds. It was used for at least one hundred years as one of the traits under selection. In a few cases, certain colours were selected against because the people at a particular time in history thought these colours typically brought health related problems with them. Indeed, some colours do. Other colours were selected against or for because the breeders felt that those colours help that breed do its job better, as in the case of the preponderance of brown coloured hunting dogs in the European hunting breeds, or as in the case of an unfortunate reddish dog who, during a hunt, was mistakenly shot for a fox, and therefore West Highland terriers are white today.

MelaninsDogs have a wide variety of genes consisting of several alleles that influence colour, and the same genes may have a very different effect on different types and lengths of coats. However, hair colour is basically determined by a substance called melanin which is produced by melanocytes which live in the skin and at the base of each hair. The more pigment granules there are, and the more tightly packed, the darker the hair. Two kinds of melanin contribute to hair color. Eumelanin is dark, though it can vary somewhat in colour due to variations in the protein that forms the framework of the pigment granule. The base form of melanin is black, but it can also appear brown or blue-grey. The second pigment, which varies from pale cream through shades of yellow, tan and red to mahogany, is called phaeomelanin. When a melanocyte switches between producing pale and dark pigments in a single hair, the effect is banded hairs, as seen in the agouti allele. When both eumelanin and pheomelanin mix together inside of one melanin granule, it’s called mixed melanin. Pigment produced in the melanocites is releases at the outer edge of the cell and is then incorporated into the shaft of the growing hair.

Looking at a hair through a videomicroscope, we can see that hair is actually translucent. Its various surfaces do not react to light in the same way, nor do they reflect it in precisely the same way. A small proportion (5 to 6%) of available light is reflected by the hair like a mirror. The rest penetrates the hair shaft, where it is absorbed to a greater or lesser extent by melanins. The melanin is housed inside granules. These melanin-filled granules are scattered through the cortex of the hair. There is no set pattern and no set amount. This is how nature creates so many variations of haircolour from only one natural substance – melanin. The color of the hair is then decided by what type of melanin is in the hair (eumelanin or phaeomelanin based) and how much melanin is in the hair and how it is distributed.

Inheritance of colour

More and more breeders and researchers of lesser known breeds with a wide variety of colour find that nature produces dogs with colours that cannot be described with the help of “traditional” genetic models (like described in the classic works of C.C. Little, O. Winge, A. G. Searle). Indeed, all of these are based on hypothesized gene variations (alleles) with hypothesized dominance order at hypothesized loci to fit data obtained from coat colours and patterns of dogs from various breeds and litters and therefore have their limitations. DNA research has shown that there are more genes involved than those hypothesized and that the actual number of alleles at genes is more for some genes and fewer for other genes than as described by classical models. Furthermore, we now know that all dogs have all genes, though in some breeds the alleles are “fixed” which means all dogs are homozygous for the same allele, and no other allele will ever appear in the course of breeding. The more coat colours occur in a breed, the more genes will be needed to explain the genotype and phenotype of the dog because no gene acts in isolation, and there are interactions among the various genes in the pathway so that some colours are not possible unless particular alleles occur at more than one locus.

Genes shaping pumi colours

Black versus red

Black versus redThe gene designated as “E” mapped to dog chromosome 5 is the receptor for Melanocyte Stimulating Hormone (MSH), which is responsible for eumelanin production. If the dominant form of E is not present, no dark pigment is produced and the dog’s coat will be reddish in colour (yellow, orange or red). On the other hand, if at least one E allele occurs, dark pigment would be the basis for a dog’s coat colour. A third allele, Em, causes reddish dogs to have a black or brown facial mask.

Brown

Blavk versus brownDifferent terms are sometimes used for this genetic colour, depending on breed and sometimes country too (i.e. brown in some breeds is known as chocolate, liver, deadgrass, cafe-au-lait etc.). The absence of the dominant form of B (Tyrosinase Related Protein 1) will change the black colour of eumelanin into brown but will have no effect on the coat colour of ee genotype dogs. E?B? genotypes will have eumelanin normally produced and will have black coat colour. E?bb genotypes will have eumelanin produced but their coat will be brown in colour. NOTE: any “bb” genotype dog will have brown nose and pad leather. The eeB? and eebb genotype dogs do not produce eumelanin pigment and will have reddish coat colour. Brown is a very rare unacceptable colour in pumis.

K Locus

Dogs have a dominant form of black completely obliterating all formation of phaeomelanin pigment. This relative newcomer to the traditional models of dog colour genetics codes both for dominant black and brindle in decreasing order of dominance, with “normal” colour at the bottom of the scale (K > Kbr > k). K is overridden by the double mutant alleles of the eumelanin producing hormone gene, so red dogs that have an ee genotype could have any combination of alleles at the K locus. A dog with at least one E and K allele (E?K?) will have an entirely eumelanin based coat and will therefore be unaffected by any other gene that has an effect on phaeomelanin. Dogs with E?kk genotype can express a variety of phenotypes, since the distribution of eumelanin and phaeomelanin in this case is determined solely by the agouti alleles present. Note: Brindling, the presence of stripes of eumelanin-based hairs in areas that are otherwise phaeomelanin based, is not known to be present in pumis.

Agouti alleles

The A gene (agouti signal peptide (ASIP), mapped to dog chromosome 24) is a bit more complicated. This gene controls where and how two pigments are produced on the body or even on the individual hair. It is very important to understand that the effects of this gene depend on the ability to produce eumelanin, which does not occur in ee genotype dogs. Therefore, ee genotype dogs remain unaffected by the agouti series.

  • The most dominant allele in this group is thought to be ay, which causes sable and fawn colouration of the coat often with hair tips dark. The extent of the eumelanin tip varies considerably from lighter sables to darker sables breed by breed. For a long time it was assumed that the differences in darkness of fawn or sable dogs are determined by whether they are homozygous or heterozygous for ay (ayay being the “clear red/yellow”, ayat being “sabled red/yellow”). It is now known that some or all of this variation is caused by another gene.
  • The next allele in dominance is aw, which is characterized by alternately banded hair and sometimes is called “wild type”. This type causes hair to change its colour during growth from light to dark resulting in bands as seen, for example, in some German Shepherds or wolves. The “competition” between ASIP and MSH is going on as the hair is growing which results in a hair that changes colour along its length.
  • Black&tan pumiThe black-and-tan allele, at, gives a black dog its tan markings around the eyes, muzzle, chest, stomach and lower legs of a primarily black dog. This is typical of many hounds as well as Rottweilers and Doberman Pinschers. The same allele is responsible for tri-colour dogs (black-and-tan with white). New studies seem to suggest another similar allele in the a series, as, for saddle, to describe the saddle tan in many terrier breeds (where black is restricted to the back and side regions, with extensive tan on the legs and head). Whether this is a separate allele or is a modification of the black&tan pattern by another gene is not yet known.
  • The last allele is a recessive black a, which is considered to be recessive to all other allele in the group. Such aa genotype dogs are black. This can be seen in German Shepherds and Shetland Sheepdogs (bi-colour).

Melanistic mask

The mechanism by which a black mask is formed is an interaction between the E gene with the agouti protein and MSH. The Em allele allows agouti to bind some of the time and cause fawn pigment to be made on the body and the melanocyte stimulating hormone to bind on the face instead. Because of this any phaeomelanin pigmented dog (i.e. yellow, fawn, red, cream) with a mask must be coloured due to an agouti genotype. Such dogs cannot be ee because Em allele is required for the production of a mask. At its weakest, the mask factor may produce black hair fringing the mouth, or a slightly smutty muzzle. At its strongest, most of the head is black, and there is considerable blackening of the chest and legs. The effect of Em shows to its fullest on clear sable dogs (ayay). In its strongest version, it can change a black&tan to a pseudo-black, with tan so restricted in its distribution that it may not be immediately apparent that the dog is not black.

Since the mask is inherited as a dominant trait, a dog could be heterozygous or homozygous for mask. The extent of the mask or depth of colour do not seem to be affected by the number of copies of em. Melanin pigment can be black, grey or brown and therefore the term melanistic mask includes all these types of masks. In some breeds all dogs have a mask, which is called fixed in genetic terms since the trait if fixed and never varies. The opposite if fixed is variable, which means some individuals in the breed have a mask and others don’t.

Some theories place the expression of a black mask at a separate locus and use the symbol Se (super-extension) for this.

Diluted or pale colours

Blue

Blue (or charcoal grey), as a dilution of black, has recently been shown to be caused by the melanophilin gene. The blue caused by this gene could be termed “born blue”, since it is present from birth. d affects both eumelanin and phaeomelanin pigment, and is thought to act by causing the clumping of pigment granules in the hair. Likeb, it often affects skin and eye colour, and in some breeds dd has been associated with skin problems and hair loss (Black Hair Follicular Dysplasia, a.k.a. Colour Dilution Alopeicia). dd is not a desirable colour in pumis, although it often occurs (silver-born and apricot pumis).

The alleles of the B and D series combine to form a range of eumelanistic (black-based) colours, out of which only those are accepted in pumis, that contain at least one B and one D allele, i.e. black colour. Blue (B?dd), liver (bbD?) and faded liver (bbdd) are not acceptable.

Progressive greying

The effect of G, in single or double dose, is the replacement of coloured by uncoloured hairs as the animal ages, very much like premature greying in human beings. This gene should be suspected in any breed where a dark puppy pales and washes out with age, and the paling is due to interspersed white colours. The fading may start immediately after birth or after a period of weeks to months has elapsed, and may go as far as it is going to by the first adult coat or may continue through the animal’s lifetime. All allele combinations of the G series are known and accepted in pumis.

Phaeomelanin dilution

This gene affects the intensity of melanin production in the coat hairs. The normal, or dominant, form is C, what might be termed “full colour”. The so-called tyrosinase (TYR) gene has been proven in mice, cattle and humans, but to date no mutations affecting dog coat colours have been found in the coding sequence of TYR.

  • The hypothetical C allele, cch (chinchilla) that is thought to lighten most or all of the phaeomelanin in the coat with little or no effect on eumelanin (e.g. causing black&tan to turn into black&silver), has not yet been found in the dog genome. If black&silver pumies are also affected by the silvering gene, the darker hairs might fade so much that the “tan” patterns will no longer be recognisable at an adult age.
  • The extreme dilution allele (ce), which is thought to cause tan to become almost white (thus affecting phaeomelanin-based coats) is also still escaping researchers.However, there certainly exists a gene that dilutes the only pigment (red/yellow) of ee dogs to cream or silver/white producing “white” dogs with full expression of dark nose and eye pigment.
  • White is often called the absence of pigment (albinism). Comb out a few hairs from your dog and lay them on a white sheet of paper. Try to decide if you see any hint of red pigment in the hairs. If so, then your dog is probably better described as cream. Tyrosinase is one gene that causes some forms of albinism in mice, people and cows. Most researchers have equated the C locus with TYR because of albino mutations in this locus. TYR has been mapped to dog chromosome 21, however, when examining the coding sequence of TYR in dogs exhibiting the classic characteristics of an albino, no mutations were found. In the same way, the alleles thought to be responsible for blue-eyed albinos (cb) and pink-eyed “real” albinos (c) have not so far been traced down in the C series. In fact, it is not known whether white coat and blue eyes in these dogs are even caused by the same gene. I am not aware of any albino pumis ever been born.

Spots and white markings

Pumi puppy with white chestmarkSo far there is no gene confirmed to cause spotting in dogs, and I do not wish
to elaborate on the traditional theories of the S series since spotting (and the connected ticking series) since they have little relevance to pumis. Pumis are self-coloured (SS), although some are known to have a tiny amount of white, especially on the chest or foot (sisi genotype). The puppy on the right was born with a few white hairs in the middle of her chest. By the time she grows up, they will have disappeared entirely. Larger spots, however, remain clearly visible. This is generally considered a fault to some degree, depending on the size of the white mark.

Standard pumi colours

- Grey in various shades (normally, the colour at birth is black, turning grey with time).
- Black.
- Fawn (fakó). Primary colours: red, yellow, cream (a trace of black or grey and a distinct mask are desirable NOTE: this does not count as clear marking).
A white mark on the chest less than 3 cm in diameter and/or a white line on the toes are not faulty.
- White.
The coat colour must always be intense and solid.

Does this pumi have a standard colour?

The following matrix helps you to decide if a certain pumi is of a standard and desirable colour, a tolerated, but not desirable colour, or an unaccepted colour.
Colour matrix

Sources and further reading

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