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This cat's coat allows it to blend in well with its environment.

Cat coat genetics affect the coloration, pattern, length, and texture is a complex subject, and many genes are involved. Cat coat genetics can produce a variety of colors and patterns. These are physical properties and should not be confused with a breed of cat. Furthermore, cats may show the color and/or pattern particular to a certain breed without actually being of that breed. For example, cats may have point coloration, but not be Siamese.

Solid colors

Eumelanin

A chocolate Oriental Shorthair kitten.

The browning gene B/b/b^l codes for TYRP1, an enzyme involved in the metabolic pathway for eumelanin pigment production. Its dominant form, B, will produce black eumelanin. It has two recessive variants, b(chocolate), and b^l(cinnamon), with b^l being recessive to both B and b.^[1] Chocolate is a rich brown color, and is referred to as chestnut in some breeds. Cinnamon is a lighter reddish brown.

This cat shows the "red" O- genotype.

Sex-linked orange/red

The sex-linked Orange gene, O/o, determines whether a cat will produce eumelanin. In cats with orange fur, phaeomelanin (red pigment) completely replaces eumelanin (black or brown pigment).^[2] This gene is located on the X chromosome. The orange allele is O, and is codominant with non-orange, o. Males can typically only be orange or non-orange due to only having one X chromosome. Since females have two X chromosomes, they have two alleles of this gene. OO results in orange fur, oo results in black or brown fur, and Oo results in a tortoiseshell cat, in which some parts of the fur are orange and others areas non-orange.^{[3][unreliable source]} Male tortoiseshell cats are known to exist, but due to the genetics involved, they almost always exhibit Klinefelter syndrome due to possessing an additional X chromosome.^{[citation needed]}

This color is known as red by breeders. Other names include yellow, ginger, and marmalade. Red show cats have a deep orange color, but it can also present as a yellow or light ginger color. Unidentified "rufousing polygenes" are theorized to be the reason for this variance.
Orange is epistatic to nonagouti,^[4] so all red cats are tabbies. "Solid" red show cats are usually low contrast ticked tabbies.

Dilution/Maltesing

The Dense pigment gene, D/d, codes for melanophilin, a protein involved in the transportation and deposition of pigment into a growing hair.^[4] When a cat has two of the recessive d alleles (Maltese dilution), black fur becomes "blue" (appearing gray), chocolate fur becomes "lilac" (appearing light brown), cinnamon fur becomes fawn, and red fur becomes cream.

A female domestic shorthair tortoiseshell cat.

Other Genes

• 
  Barrington Brown is a recessive browning gene that dilutes black to mahogany, brown to light brown and chocolate to pale coffee. It is different from the browning gene and has only been observed in laboratory cats.^[5]
  


• The Dilution modifier gene, Dm, "caramelizes" the dilute colors as a dominant trait. The existence of this phenomenon as a discrete gene is a controversial subject among feline enthusiasts.


• A mutation at the extension locus E/e (the melanocortin receptor) changes black pigment to amber or light amber. This phenomenon was first identified in Norwegian Forest cats.^[6]
  


• A modifying factor has also been hypothesized in shaded silver and chinchilla Persians whose fur turns pale golden in adulthood, due to low levels of phaeomelanin production. These cats resemble shaded or tipped goldens, but are genetically shaded or tipped silvers. This is probably related to the phenomenon known as "tarnishing" in silvers.

Tabbies

A mackerel tabby with the classic "M" on forehead.

Tabby cats are striped due to the agouti gene. Their stripes have an even distribution of pigment, while the background is made up of banded hairs. Tabby cats usually show the following traits:

• M on forehead. (Visible in ticked tabby cats, but hard to discern in shaded silver/golden, and tipped cats)


• Thin pencil lines on face. (Visible in ticked tabby cats, but hard to discern in shaded silver/golden, and tipped cats)


• Black "eyeliner" appearance and white or pale fur around eyeliner.


• Pigmented lips and paws.


• A pink nose outlined in darker pigment.


• Torso, leg, and tail banding. (Torso banding disappears in the ticked tabby.)

Agouti

An Abyssinian kitten displaying the ticked tabby pattern.

The Agouti gene, A/a, codes for agouti signaling protein. The dominant, wild-type A causes the agouti shift phenomenon which causes hairs to be banded with black and red (revealing the underlying tabby pattern), while the recessive non-agouti or "hypermelanistic" allele, a, prevents this shift in the pigmentation pathway. In its homozygous form, aa, this results in black pigment production throughout the growth cycle of the hair.^[7] Thus, the non-agouti genotype (aa) masks or hides the tabby pattern, although sometimes a suggestion of the underlying pattern can be seen (called "ghost striping"), especially in kittens. The O allele is also epistatic over the aa genotype. That is, the A to a mutation does not have a discernible effect on red or cream colored cats, resulting in these cats displaying tabby striping independent of their genotype at this locus. This explains why you can usually see the tabby pattern in the orange patches of non-agouti tortoiseshell cats, but not in the black or brown patches.

Tabby Pattern

Down, awn and guard hairs of a domestic tabby cat

The primary Tabby gene, Mc/mc, sets the basic pattern of stripes that underlies the coat. The basic wild-type tabby gene, Mc, produces what is called a mackerel striped tabby (stripes look like thin fishbones and may break up into bars or spots), while a recessive mutant, mc, produces a blotched or classic tabby pattern (broad bands, whorls, and spirals of dark color on pale background usually with bulls-eye or oyster pattern on flank). The classic tabby is most common in Iran, Great Britain and in lands that were once part of the British Empire and Persian Empire. The gene responsible for this differential patterning has been identified as transmembrane aminopeptidase Q (Taqpep), which also produces the king cheetah coat variant.^[8]

Spotted Tabby

Spotted tabbies have their stripes broken up into spots, which may be arranged vertically or horizontally. A 2010 study suggests that spotted is caused by the modification of mackerel stripes, and may cause varying phenotypes such as "broken mackerel" tabbies via multiple loci.^[9]

Ticked Tabby

The Ticked gene, T^a / t^a, produces an Abyssinian ticked or non-patterned agouti tabby with the dominant allele, having virtually no stripes or bars. Stripes usually remain to some extent on the face, tail, legs, and sometimes the chest. The dominant ticked tabby allele masks all other tabby patterns.^[9]

Other Genes

• Other genes (pattern modifier genes) are theorized to be responsible for creating various type of spotting patterns, many of which are variations on a basic mackerel or classic pattern. There are also hypothetical factors which affect the timing and frequency of the agouti shift, affecting agouti band width and the number and quality of alternating bands of eumelanin and phaeomelanin on individual hairs.


• There is a gene, not yet identified but believed to be related to the agouti gene, in the Chausie breed, that produces silver-tipped black fur similar to Abyssinian ticked fur. The "grizzled" phenomenon is purported to have been inherited from the hybridization of the domestic cat to the jungle cat (Felis chaus).


• The inhibited pigment gene, I/i. The dominant allele (I) produces tipped hairs that are fully colored only at the tip and have a white base. This allele appears to interact with other genes to produce various degrees of tipping, ranging from deeply tipped silver tabby to lightly tipped shaded silver and chinchilla silver. The inhibitor gene interacts with the non-agouti genotype (I-aa) to produce the color known as smoke. The homozygous recessive genotype when combined with the agouti gene (iiA-), produces tabby coloration, which can vary along a spectrum ranging from a deeply patterned brown tabby, to a lighter "golden tabby", to the very lightly colored shaded or chinchilla golden colors. Orange cats with the inhibitor gene (I-O-) are commonly called "cameo".

Tortoiseshells and calicos

Female tortoiseshell-and-white cat

Tortoiseshells are also known as calimanco or clouded tiger cats, and by the abbreviation "tortie". Tortoiseshells have patches of orange fur (pheomelanin based) and black or brown (eumelanin based) fur, caused by X-inactivation. Because this requires two X chromosomes, the vast majority of tortoiseshells are female, with approximately 1 in 3,000 being male.^[10] Male tortoiseshells can occur as a result of chromosomal abnormalities such as Klinefelter syndrome, or by a phenomenon known as chimerism, where two early stage embryos are merged into a single kitten.

Tortoiseshells with a relatively small amount of white spotting are known as "tortoiseshell and white", while those with a larger amount are known in North America as calicos. Calicos are also known as tricolor cats, mi-ke (meaning "triple fur") in Japanese, and lapjeskat (meaning "patches cat") in Dutch. The factor that distinguishes tortoiseshell from calico is the pattern of eumelanin and pheomelanin, which is partly dependent on the amount of white, due to an effect of the white spotting gene on the general distribution of melanin. A cat which has both an orange and non-orange gene, Oo, and little to no white spotting, will present with a mottled blend of red/cream and black/blue, reminiscent of tortoiseshell material, and is called a tortoiseshell cat. An Oo cat with a large amount of white will have bigger, clearly defined patches of red/cream and black/blue, and is called a calico. With intermediate amounts of white, a cat may exhibit a calico pattern, a tortie pattern, or something in between, depending on other epigenetic factors.

A blue calico British Shorthair.

A true tricolor must consist of three colors: white; a red, orange, yellow, or cream pheomelanin color; and a black, brownish, or gray (blue) eumelanin color. Tricolor should not be mistaken for the natural gradations in a tabby pattern. The shades which are present in the pale bands of a tabby are not considered to constitute a separate color.^[11]

Variations

• The basic tortoiseshell pattern has several different colors depending on the color of the eumelanin (the B locus), and dilution (the D locus).


• Tortoiseshell tabbies, also known as torbies, display tabby patterning on both colors. Calico tabbies are also called calibys.^[12]
  

White spotting and epistatic white

Blue (grey) and white bicolor cat

White spotting and epistatic white (also known as dominant white) were long thought to be two separate genes, but in fact they are both on the KIT gene. White spotting can take many forms, from a small spot of white to the mostly-white pattern of the Turkish Van, while epistatic white produces a fully white cat. The Birman-specific recessive "gloving" trait is also located on the KIT gene.

• W^D = dominant white, linked to blue eyes and deafness. The deafness is due to a reduction in the population and survival of melanoblast stem cells, which in addition to creating pigment-producing cells, develop into a variety of neurological cell types. White cats with one or two blue eyes have a particularly high likelihood of being deaf.


• W^S = white spotting. It exhibits codominance and variable expression; heterozygous cats have somewhere between 0-50% white, and homozygous cats have between 50-100% white.


• w = wild type, no white spotting.


• w^g = recessive Birman white gloving allele.

Colorpoint and albinism

Siamese cat, classical colorpoint pattern

The colorpoint pattern is most commonly associated with Siamese cats, but may also appear in any domesticated cat. A colorpointed cat has dark colors on the face, ears, feet, and tail, with a lighter version of the same color on the rest of the body, and possibly some white. The exact name of the colorpoint pattern depends on the actual color, so there are seal points (dark brown), chocolate points (warm lighter brown), blue points (dark gray), lilac or frost points (silvery gray-pink), red or flame points (orange), and tortie (tortoiseshell mottling) points, among others. This pattern is the result of a temperature sensitive mutation in one of the enzymes in the metabolic pathway from tyrosine to pigment, such as melanin; thus, little or no pigment is produced except in the extremities or points where the skin is slightly cooler. For this reason, colorpointed cats tend to darken with age as bodily temperature drops; also, the fur over a significant injury may sometimes darken or lighten as a result of temperature change.

The tyrosine pathway also produces neurotransmitters, thus mutations in the early parts of that pathway may affect not only pigment, but also neurological development. This results in a higher frequency of cross-eyes among colorpointed cats, as well as the high frequency of cross-eyes in white tigers.

• C = full color.


• cb = Burmese "sepia" pattern, similar to colorpoint but with lower contrast.


• cs = Siamese/colorpoint. It is codominant with cb; cb/cs cats show a medium-contrast phenotype known as mink.


• ca = Blue-eyed albino.


• c = Pink-eyed albino.

Silver and golden series

A silver tabby British Shorthair.

The silver series is caused by the Melanin inhibitor gene I/i. The dominant form causes melanin production to be suppressed, but it affects phaeomelanin (red pigment) much more than eumelanin (black or brown pigment). On tabbies, this turns the background a sparkling silver color while leaving the stripe color intact, making a silver tabby. On solid cats, it turns the base of the hair pale, making them silver smoke.^[13]

Silver agouti cats can have a range of phenotypes, from silver tabby, to silver shaded (under half the hair is pigmented), to tipped silver/chinchilla (only the very tip of the hair is pigmented). This seems to be affected by hypothetical wide band factors, which make the silver band at the base of the hair wider. Breeders often notate wide band as a single gene Wb/wb, but it is most likely a polygenic trait.

If a cat has the wide band trait but no inhibitor, the band will be golden instead of silver. These cats are known as golden tabbies. Shaded golden and tipped golden are also possible. However, there is no golden smoke, because the combination of wide band and nonagouti simply produces a solid cat.^{[14][unreliable source?]}

A golden tabby Siberian cat.

The genetics involved in producing the ideal tabby, tipped, shaded, or smoke cat is complex. Not only are there many interacting genes, but genes sometimes do not express themselves fully, or conflict with one another. For example, the melanin inhibitor gene in some instances does not block pigment, resulting in a grayer undercoat, or in tarnishing (yellowish or rusty fur). The greyer undercoat is less desirable to fanciers.

Likewise, poorly-expressed non-agouti or over-expression of melanin inhibitor will cause a pale, washed out black smoke. Various polygenes (sets of related genes), epigenetic factors, or modifier genes, as yet unidentified, are believed to result in different phenotypes of coloration, some deemed more desirable than others by fanciers.

Tipped or shaded cats

Here are the genetic influences on tipped or shaded cats:

• 
  Agouti gene.


• Tabby pattern genes (such as T^a masking the tabby pattern).


• Silver/melanin inhibitor gene.


• Factors affecting the number and width of bands of color on each hair (such as the hypothetical wide band gene).


• Factors affecting the amount and quality of eumelanin and/or phaeomelanin pigment expression (such as theorized rufousing factors)


• Genes causing sparkling appearance (such as glitter in the Bengal, satin in the Tennessee Rex, grizzle in the Chausie).


• Factors to clear up residual striping (hypothetical Chaos, Confusion, Unconfused, Erase, and Roan factors).

Fever coat

Fever coat is an effect known in domestic cats, where a pregnant female cat has a fever or is stressed, causing her unborn kittens' fur to develop a silver-type color (silver-grey, cream, or reddish) and not what the kitten's genetics would cause. After birth, over some weeks the silver fur is replaced naturally by fur colored according to the kitten's genetics.^[15][16][17]

Fur length and texture

A Devon Rex.

Cat fur length is governed by the Long hair gene in which the dominant form, L, codes for short hair, and the recessive l codes for long hair. In the longhaired cat, the transition from anagen (hair growth) to catagen (cessation of hair growth) is delayed due to this mutation.^[18] A rare recessive shorthair gene has been observed in some lines of Persian cat (silvers) where two longhaired parents have produced shorthaired offspring.

There have been many genes identified that result in unusual cat fur. These genes were discovered in random-bred cats and selected for. Some of the genes are in danger of going extinct because the cats are not sold beyond the region where the mutation originated or there is simply not enough demand for cats expressing the mutation.

In many breeds, coat gene mutations are unwelcome. An example is the rex allele which appeared in Maine Coons in the early 1990s. Rexes appeared in America, Germany and the UK, where one breeder caused consternation by calling them "Maine Waves". Two UK breeders did test mating which indicated that this was probably a new rex mutation and that it was recessive. The density of the hair was similar to normally coated Maine Coons, but consisted only of down type hairs with a normal down type helical curl, which varied as in normal down hairs. Whiskers were more curved, but not curly. Maine Coons do not have awn hairs, and after moulting, the rexes had a very thin coat.

Curly-coated

There are various genes producing curly-coated or "rex" cats. New types of rex arise spontaneously in random-bred cats now and then. Here are some of the rex genes that breeders have selected for:

• r = Cornish Rex, recessive.


• gr (provisional) = German Rex, recessive. Same locus as Cornish, but proposed as a different allele. However, most breeders consider the German Rex to have r/r genotype.


• re = Devon Rex, recessive.


• ro = Oregon Rex (extinct), recessive.


• Se = Selkirk Rex, dominant although sometimes described as an incomplete dominant because the three possible allele pairings relate to three different phenotypes: heterozygous cats (Sese) may have a fuller coat that is preferred in the show ring, while homozygous cats (SeSe) may have a tighter curl and less coat volume. (sese type cats have a normal coat.) This phenomenon may also colloquially be referred to as additive dominance.


• Lp (provisional) = LaPerm, dominant: Lp/lp and Lp/Lp individuals have the same phenotype.

Hairlessness

There are also genes for hairlessness:

• h = French hairless cat, recessive.


• hd = British hairless cat, recessive.


• Hp = Russian Donskoy and Peterbald, dominant.


• hr = Canadian Sphynx cat, recessive.

Some rex cats are prone to temporary hairlessness, known as baldness, during moulting.

Here are a few other genes resulting in unusual fur:

• The Wh gene (dominant, possibly incomplete) results in Wirehair cats. They have bent or crooked hair producing springy, crinkled fur.


• A hypothetical Yuc gene, or York Chocolate undercoat gene, results in cats with no undercoat. However, the proportional relationship between guard, awn, and down hair production varies greatly between all breeds.


• A recessive autosomal gene for Onion hair which causes roughness and swelling on the hairs. The swelling is due to enlargement of the inner core of medulla cells.


• A recessive autosomal gene spf for sparse fur. As well as sparse coat, the hairs are thin, straggly and contorted and there is brown exudate around the eyes and nose and on the chest and stomach. A similar condition is linked to Ornithine Transcarbamylase Deficiency in mice.

See also

• Bicolor cat


• Calico cat


• Tortoiseshell cat


• Point (coat color)


• Barr body


• Tabby cat


• Maltese cat


• Deaf white cat


• Dog coat genetics


• Equine coat color genetics

References

Further reading

• 
  Vella, Carolyn M.; Shelton, Lorraine M.; McGonagle, John J.; Stanglein, Terry W. (1999). Genetics for Cat Breeders and Veterinarians. Butterworth-Heinemann Ltd. ISBN 0-7506-4069-3.
  


• 
  Jude, A. C. (1955). Cat Genetics.
  


• 
  Lorimer, Heather E. "Coat Color Genetics".
  

External links

• Cat Genetics articles in plain English


• Cat Gene Symbols


• The Pigment Parade: White and White Spotting in the Cat


• Inhibitor gene diagram and article