Cat Coat Genetics
   HOME

TheInfoList



OR:

Cat coat genetics determine the coloration, pattern, length, and texture of feline fur. The variations among cat coats are physical properties and should not be confused with cat breeds. A cat may display the coat of a certain breed without actually being that breed. For example, a Neva Masquerade (
Siberian Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states si ...
colorpoint) could wear
point coloration Points are specific areas of an animal coat that are colored differently from the main body colorations. Point coloration may be represented by a pale body color and relatively darker extremities, such as face, ears, feet, tail, and external sex ...
, the stereotypical coat of a Siamese.


Solid colors


Eumelanin and phaeomelanin


Eumelanin

The browning gene ''B/b/bl'' codes for
TYRP1 Tyrosinase-related protein 1, also known as TYRP1, is an intermembrane enzyme which in humans is encoded by the ''TYRP1'' gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is ...
(), 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 ''bl'' (cinnamon), with ''bl'' being recessive to both ''B'' and ''b''. Chocolate is a rich dark brown color, and is referred to as chestnut in some breeds. Cinnamon is a light brown which may be a reddish color.


Sex-linked red

The
sex-linked Sex linkage describes the sex-specific patterns of inheritance and expression when a gene is present on a sex chromosome (allosome) rather than a non-sex chromosome (autosome). Genes situated on the X-chromosome are thus termed X-linked, and a ...
red ''"Orange"'' locus, ''O/o'', determines whether a cat will produce eumelanin. In cats with orange fur,
phaeomelanin Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...
(red pigment) completely replaces
eumelanin Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...
(black or brown pigment). This gene is located on the
X chromosome The X chromosome is one of the two sex chromosomes in many organisms, including mammals, and is found in both males and females. It is a part of the XY sex-determination system and XO sex-determination system. The X chromosome was named for its u ...
. The orange allele is O, and non-orange is o. Males are typically only 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 fur without any orange (black, brown, etc.), and Oo results in a
tortoiseshell cat Tortoiseshell is a cat animal coloration, coat coloring named for its similarity to tortoiseshell pattern. Like tortoiseshell-and-white or Calico cat, calico cats, tortoiseshell cats are almost exclusively female. Male tortoiseshells are rare and ...
, in which some parts of the fur are orange and other areas non-orange. One in three thousand tortoiseshell cats are male, making the combination possible but rare - however, due to the nature of their genetics, male tortoiseshells often exhibit chromosomal abnormalities. In one study, less than a third of male calicos had a simple XXY Klinefelter's karyotype, slightly more than a third were complicated XXY mosaics, and about a third had no XXY component at all. The pelt color commonly referred to as "orange" is scientifically known as red. Other common 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
polygene A polygene is a member of a group of non- epistatic genes that interact additively to influence a phenotypic trait, thus contributing to multiple-gene inheritance (polygenic inheritance, multigenic inheritance, quantitative inheritance), a type o ...
s" are theorized to be the reason for this variance. ''Orange'' is
epistatic Epistasis is a phenomenon in genetics in which the effect of a gene mutation is dependent on the presence or absence of mutations in one or more other genes, respectively termed modifier genes. In other words, the effect of the mutation is depe ...
to ''nonagouti'', so all red cats are tabbies. "Solid" red show cats are usually low contrast ticked tabbies. The identity of the gene at the ''Orange'' locus was narrowed down to a 3.5 Mb stretch on the X chromosome in 2009. In 2024 it was discovered that the dominant orange color associated with the ''Orange'' locus is the result of a genomic deletion in a regulatory region o
Arhgap36
a Rho GTPase activating protein. The deletion results in a 13-fold increase in expression of the protein in melanocytes.


Dilution

The ''Dense pigment'' gene, ''D/d'', codes for melanophilin (''MLPH''; ), a protein involved in the transportation and deposition of pigment into a growing hair. When a cat has two of the recessive ''d'' alleles (Maltese dilution), black fur becomes "blue" (appearing gray), chocolate fur becomes "lilac" (appearing light, almost grayish brown-lavender), cinnamon fur becomes "fawn", and red fur becomes "cream". Similar to red cats, all cream cats are tabbies. The ''d'' allele is a single-base deletion that truncates the protein. If the cat has d/d genes, the coat is diluted. If the genes are D/D or D/d, the coat will be unaffected.


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. * 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. * ''Amber'', a mutation at the extension locus ''E/e'' (the
melanocortin 1 receptor The melanocortin 1 receptor (MC1R), also known as melanocyte-stimulating hormone receptor (MSHR), melanin-activating peptide receptor, or melanotropin receptor, is a G protein–coupled receptor that binds to a class of pituitary peptide hormon ...
, ''MC1R'') changes black pigment to amber or light amber, similar in appearance to red and cream. Kittens are born dark but lighten up as they age. Paws and nose still exhibit the original undiluted color in contrast to other diluted colors, where paws and nose have the diluted color. This phenomenon was first identified in Norwegian Forest cats. * Another recessive mutation at extension was discovered which causes the russet color in Burmese cats. It is symbolized as er. Like amber cats, russet cats lighten as they age. * A modifying factor has also been hypothesized in shaded silver and chinchilla
Persians Persians ( ), or the Persian people (), are an Iranian ethnic group from West Asia that came from an earlier group called the Proto-Iranians, which likely split from the Indo-Iranians in 1800 BCE from either Afghanistan or Central Asia. They ...
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

Tabby cats have a range of variegated and blotched coats, consisting of a dark pattern on a lighter background. This variety is derived from the interplay of multiple genes and resulting phenotypes. Most tabbies feature thin dark markings on the face, including the 'M' on the forehead and an eyeliner effect, pigmented lips and paws, and a pink nose outlined in darker pigment. However, the following different coat patterns are all possible: * Mackerel: Thin, dark stripes (sometimes called "tiger stripes") * Blotched/Classic: Thicker bands or whorls of dark pigment. * Spotted: Broken bands that look more like individual spots. * Ticked: No distinct stripes, spots, or blotches on the body—though some may be visible on the legs, face, and tail. File:野良猫のキジトラ (2015 photo; cropped 2022).jpg, Mackerel File:MaineCoonSansFond.png, Blotched / classic File:Star Spangled Cat (mirrored).jpg, Spotted File:Abyssin.jpg, Ticked


Agouti

The agouti factor determines the "background" of the tabby coat, which consists of hairs that are banded with dark eumelanin and lighter phaeomelanin along the length of the hair shaft. The ''Agouti'' gene, with its dominant ''A'' allele and recessive ''a'' allele, controls the coding for agouti signaling protein (ASIP; ). The wild-type dominant ''A'' causes the banding and thus an overall lightening effect on the hair, while the recessive ''non-agouti'' or "hypermelanistic" allele ''a'' does not initiate this shift in the pigmentation pathway. As a result, homozygous ''aa'' have pigment production throughout the entire growth cycle of the hair and therefore along its full length. These homozygotes are solidly dark throughout, which obscures the appearance of the characteristic dark tabby markings—sometimes a suggestion of the underlying pattern, called "ghost striping", can be seen, especially in bright slanted light on kittens and on the legs, tail and sometimes elsewhere on adults. A major exception to the solid masking of the tabby pattern exists, as the ''O'' allele of the ''O/o'' locus is
epistatic Epistasis is a phenomenon in genetics in which the effect of a gene mutation is dependent on the presence or absence of mutations in one or more other genes, respectively termed modifier genes. In other words, the effect of the mutation is depe ...
over the ''aa'' genotype. That is, in red or cream colored cats, tabby marking is displayed regardless of the genotype at the agouti locus. However, some red and most cream tabbies do have a fainter pattern when lacking an agouti allele, indicating that the ''aa'' genotype does still have a faint effect even if it does not induce complete masking. The mechanism of this process is unknown. An example of the ''Agouti'' gene can be seen in Bengal cats, which are a hybrid between Asian Leopard cats and domestic cats. The breed has a characteristically dark face marking and a stripe down its back. This is term as a
"mask"
an
"cape"
as well as charcoal markings, according to Gershoney et. al. The charcoal mask is indicated to be the result of a heterozygote of
APbe/a.
he relationships between the different a''gouti'' haplotypes is not fully understood. More research to required to determine "modes of inheritance for charcoal" in Bengal cats.


Dark markings

The ''Tabby'' locus on chromosome A1 accounts for most tabby patterns seen in domestic cats, including those patterns seen in most breeds. The dominant allele ''TaM'' produces mackerel tabbies, and the recessive ''Tab'' produce classic (sometimes referred to as blotched) tabbies. The gene responsible for this differential patterning has been identified as ''transmembrane aminopeptidase Q'' ('' Taqpep'', ). A
threonine Threonine (symbol Thr or T) is an amino acid that is used in the biosynthesis of proteins. It contains an α-amino group (which is in the protonated −NH form when dissolved in water), a carboxyl group (which is in the deprotonated −COO− ...
to
asparagine Asparagine (symbol Asn or N) is an α-amino acid that is used in the biosynthesis of proteins. It contains an α-amino group (which is in the protonated −NH form under biological conditions), an α-carboxylic acid group (which is in the depro ...
substitution at residue 139 ( T139N) in this protein is responsible for producing the tabby phenotype in domestic cats. In
cheetah The cheetah (''Acinonyx jubatus'') is a large Felidae, cat and the Fastest animals, fastest land animal. It has a tawny to creamy white or pale buff fur that is marked with evenly spaced, solid black spots. The head is small and rounded, wit ...
s, a base pair insertion into exon 20 of the protein replaces the 16 C-terminal residues with 109 new ones (N977Kfs110), generating the king cheetah coat variant. The wild-type (in
African wildcat The African wildcat (''Felis lybica'') is a small wildcat species with sandy grey fur, pale vertical stripes on the sides and around the face. It is native to Africa, West and Central Asia, and is distributed to Rajasthan in India and Xinjiang ...
s) is the mackerel tabby (stripes look like thin fishbones and may break up into bars or spots). The most common variant is the ''classic'' tabby pattern (broad bands, whorls, and spirals of dark color on pale background usually with bulls-eye or oyster pattern on flank). Spotted tabbies have their stripes broken up into spots, which may be arranged vertically or horizontally. A 2010 study suggests that spotted coats are caused by the modification of mackerel stripes, and may cause varying phenotypes such as "broken mackerel" tabbies via multiple loci. If the genes are Sp/Sp or Sp/sp the tabby coat will be spotted or broken. If it is an sp/sp gene, the tabby pattern will remain either mackerel or blotched. This gene has no effect on cats with a ticked coat.


Ticked tabby

The ''Ticked'' (''Ti'') locus on chromosome B1 controls the generation of "ticked coats", agouti coats with virtually no stripes or bars. Ticked tabbies are rare in the random-bred population, but fixed in certain breeds such as the Abyssinian and Singapura. ''TiA'' is the dominant allele that produces ticked coats; ''Ti+'' is the recessive one. The causative gene for ticked tabby markings is ''Dickkopf-related protein 4'' ('' DKK4''). Both a
cysteine Cysteine (; symbol Cys or C) is a semiessential proteinogenic amino acid with the chemical formula, formula . The thiol side chain in cysteine enables the formation of Disulfide, disulfide bonds, and often participates in enzymatic reactions as ...
to
tyrosine -Tyrosine or tyrosine (symbol Tyr or Y) or 4-hydroxyphenylalanine is one of the 20 standard amino acids that are used by cells to synthesize proteins. It is a conditionally essential amino acid with a polar side group. The word "tyrosine" is ...
substitution at residue 63 ( C63Y) and an
alanine Alanine (symbol Ala or A), or α-alanine, is an α-amino acid that is used in the biosynthesis of proteins. It contains an amine group and a carboxylic acid group, both attached to the central carbon atom which also carries a methyl group sid ...
to
valine Valine (symbol Val or V) is an α-amino acid that is used in the biosynthesis of proteins. It contains an α- amino group (which is in the protonated −NH3+ form under biological conditions), an α- carboxylic acid group (which is in the deproton ...
substitution at residue 18 ( A18V) result in decreased ''DKK4'', which is associated with ticking. Both variants are present in the Abyssinian breed, and the A18V variant is found in the Burmese breed. Stripes often remain to some extent on the face, tail, legs, and sometimes the chest ("bleeding through"). Traditionally, this has been thought to happen in heterozygotes (''TiATi+'') but be nearly or completely nonexistent in homozygotes (''TiATiA''). The ticked tabby allele is epistatic to and therefore completely (or mostly) masks all the other tabby alleles, "hiding" the patterns they would otherwise express. It was once thought that ''TiA'' was an allele of the ''Tabby'' gene, called ''Ta'', dominant to all other alleles at the locus.


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 The Chausie () is a domestic breed of cat that was developed by breeding a few individuals from the non-domestic species jungle cat ('' Felis chaus'') to a far greater number of domestic cats ('' Felis catus''). The Chausie was first recognized ...
breed that produces silver-tipped black fur similar to Abyssinian ticked fur, known as "grizzled". This phenomenon is purported to have been inherited from the hybridization of the domestic cat to the
jungle cat The jungle cat (''Felis chaus''), also called reed cat and swamp cat, is a medium-sized cat native from the Eastern Mediterranean region and the Caucasus to parts of Central, South and Southeast Asia. It inhabits foremost wetlands like swamps, ...
(''Felis chaus''). * The rosette tabby pattern is a pattern similar to that of a
leopard The leopard (''Panthera pardus'') is one of the five extant cat species in the genus ''Panthera''. It has a pale yellowish to dark golden fur with dark spots grouped in rosettes. Its body is slender and muscular reaching a length of with a ...
, where rosette spots are spread over the body. The pattern is found in
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two diff ...
cat breeds, such as the
Bengal Bengal ( ) is a Historical geography, historical geographical, ethnolinguistic and cultural term referring to a region in the Eastern South Asia, eastern part of the Indian subcontinent at the apex of the Bay of Bengal. The region of Benga ...
and
Safari A safari (; originally ) is an overland journey to observe wildlife, wild animals, especially in East Africa. The so-called big five game, "Big Five" game animals of Africa – lion, African leopard, leopard, rhinoceros, African elephant, elep ...
. * 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 Smoke is an aerosol (a suspension of airborne particulates and gases) emitted when a material undergoes combustion or pyrolysis, together with the quantity of air that is entrained or otherwise mixed into the mass. It is commonly an unwante ...
. 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. Red and cream cats with the inhibitor gene (I-O-) are commonly called "cameo".


Tortoiseshells and calicos

Tortoiseshells have patches of orange fur (pheomelanin based) and black or brown (eumelanin based) fur, caused by
X-inactivation X-inactivation (also called Lyonization, after English geneticist Mary Lyon) is a process by which one of the copies of the X chromosome is inactivated in therian female mammals. The inactive X chromosome is silenced by being packaged into ...
. Because this requires two X chromosomes, the vast majority of tortoiseshells are female, with approximately 1 in 3,000 being male. Male tortoiseshells can occur as a result of chromosomal abnormalities such as
Klinefelter syndrome Klinefelter syndrome (KS), also known as 47,XXY, is a chromosome anomaly where a male has an extra X chromosome. These complications commonly include infertility and small, poorly functioning testicles (if present). These symptoms are often n ...
, by
mosaicism Mosaicism or genetic mosaicism is a condition in which a multicellular organism possesses more than one genetic line as the result of genetic mutation. This means that various genetic lines resulted from a single fertilized egg. Mosaicism is o ...
, 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 the United States 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 black/red and blue/cream, reminiscent of
tortoiseshell Tortoiseshell or tortoise shell is a material produced from the shells of the larger species of tortoise and turtle, mainly the hawksbill sea turtle, which is a critically endangered species according to the IUCN Red List largely because of its ...
material, and is called a tortoiseshell cat. An Oo cat with a large amount of white will have bigger, clearly defined patches of black/red and blue/cream, and is called a calico in the US. With intermediate amounts of white, a cat may exhibit a calico pattern, a tortie pattern, or something in between, depending on other epigenetic factors. Blue tortoiseshell, or ''diluted calico'', cats have a lighter coloration (blue/cream) and are sometimes called ''calimanco'' or ''clouded tiger''. A true tricolor must consist of three colors: white, a red-based color like ginger or cream, and black-based color like black or blue. 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.


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 red- and black-based colors. Calico tabbies are also called ''calibys'' or ''tabicos''. Tortoiseshell cats with small white patches are called ''tortico'' cats, a portmanteau of ''calico'' and ''tortoiseshell''.


White spotting and epistatic white

The KIT gene determines whether or not there will be any white in the coat, except when a solid white coat is caused by albinism. White spotting and epistatic white (also known as dominant white) were long thought to be two separate genes (called ''S'' and ''W'' respectively), but in fact they are both on the KIT gene. The two have been combined into a single ''white spotting'' locus (''W''). 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 (solid or self white). The KIT gene ''W'' locus has the following alleles: * WD (or W)=dominant white (solid/self white), autosomal dominant allele. It causes complete white coloration by disrupting replication and migration of
melanocyte Melanocytes are melanin-producing neural-crest, neural crest-derived cell (biology), cells located in the bottom layer (the stratum basale) of the skin's epidermis (skin), epidermis, the middle layer of the eye (the uvea), the inner ear, vagina ...
s into the skin. The carriers of this allele are white regardless of any other color-associated gene. It is linked to blue eyes and congenital sensorineural 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. Dominant white is distinct from albinism (''c'') which results from a mutation in a different gene that has no known impact on hearing. * wS (or S)=white spotting ( bicolor or tricolor cats), dominant allele. It only disrupts migration of
melanocyte Melanocytes are melanin-producing neural-crest, neural crest-derived cell (biology), cells located in the bottom layer (the stratum basale) of the skin's epidermis (skin), epidermis, the middle layer of the eye (the uvea), the inner ear, vagina ...
s to certain patches in the skin, thus leading to the formation of white spots. It exhibits codominance and variable expression: ** heterozygote (Wh or ''Ss'')= high degree of spotting white (between 0–50% white); bicolor/tricolor or ventral white (usually the feet, nose, chest, and belly), which is dominant to solid color. Heterozygous cats have somewhere between 0-50% white. ** homozygote (Wl or ''SS'')=low degree of spotting white (between 50 and 100% white); dominant harlequin and van pattern. The van pattern is named after the
Lake Van Lake Van (; ; ) is the largest lake in Turkey. It lies in the Eastern Anatolia Region of Turkey in the provinces of Van Province, Van and Bitlis Province, Bitlis, in the Armenian highlands. It is a Salt lake, saline Soda lake, soda lake, receiv ...
region in Turkey, and expresses as coloration limited to the head and tail. * w (or N)=wild-type or normal (non-white coats), recessive allele. Homozygotes for it won't have any white in their coat. * wg=Birman white gloving allele, recessive allele. * wsal=salmiak allele, recessive allele. In addition to a white tuxedo pattern, hairs are colored at the root fading into white at the tips, except on the head, where they are often colored all the way through. It was found in Finland and the name means "salty licorice".


Colorpoint and albinism

The colorpoint pattern is most commonly associated with Siamese cats, but due to crossbreeding may also appear in any (non-pedigree) domesticated cat. A colorpoint 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. A few examples are seal points (dark brown to black), chocolate points (warm, lighter brown), blue points (gray), lilac or frost points (silvery gray-pink), red or flame points (orange), and tortie (tortoiseshell mottling) points. This pattern is the result of a temperature sensitive mutation in one of the
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s in the
metabolic pathway In biochemistry, a metabolic pathway is a linked series of chemical reactions occurring within a cell (biology), cell. The reactants, products, and Metabolic intermediate, intermediates of an enzymatic reaction are known as metabolites, which are ...
from
tyrosine -Tyrosine or tyrosine (symbol Tyr or Y) or 4-hydroxyphenylalanine is one of the 20 standard amino acids that are used by cells to synthesize proteins. It is a conditionally essential amino acid with a polar side group. The word "tyrosine" is ...
to pigment, such as
melanin Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...
; thus, little or no pigment is produced except in the extremities or points where the skin is slightly cooler. For this reason, colorpoint 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. More specifically, the ''albino'' locus contains the gene '' TYR'' (). Two distinct alleles causing blue-eyed and pink-eyed albinism respectively have been previously theorized. Although the Siamese colorpoint pattern is the most famous coloration produced by ''TYR'', there are color mutations at the locus. * ''C'' is the wildtype allele resulting in full pigmentation and is completely dominant to all other known alleles at the locus. * Point=''cs'' is the ''point'' allele associated with the Siamese colorpoint pattern. * Sepia=''cb'' is an allele called ''sepia'' (or solid), and is most associated with Burmese cats. It produces a pattern similar to the Siamese colorpoint, but with a much lower contrast and amber-yellow to green eyes. * Mink=''cs'' and ''cb'' are codominant, with ''cb''/''cs'' cats having an intermediate phenotype termed ''mink'', in which the pigment distribution is between sepia and point, and the eye color is blue-green ( aquamarine). * Albinism=''c'' and c2 are two synonymous alleles recessive to all other alleles at the locus that cause albinism. * ''cm'' is a novel mutation in Burmese cats that results in a color pattern named mocha. Its interactions with other alleles have not yet been fully established. The tyrosine pathway also produces
neurotransmitter A neurotransmitter is a signaling molecule secreted by a neuron to affect another cell across a Chemical synapse, synapse. The cell receiving the signal, or target cell, may be another neuron, but could also be a gland or muscle cell. Neurotra ...
s, 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 colorpoint cats, as well as the high frequency of cross-eyes in white tigers. File:Siamese cat Vaillante.JPG, Black point Siamese File:Brown-mink.JPG, Black mink Tonkinese File:Geisha One Year Old (5640618397).jpg, Black sepia Burmese File:DSH blotched transparent (cropped).png, Black blotched
tabby A tabby cat, or simply tabby, is any domestic cat (''Felis catus'') with a coat pattern distinguished by an M-shaped marking on its forehead, stripes by its eyes and across its cheeks, along its back, around its legs and tail, and characteris ...
(lynx) point cat File:Tortie-point.jpg, Black
tortoiseshell Tortoiseshell or tortoise shell is a material produced from the shells of the larger species of tortoise and turtle, mainly the hawksbill sea turtle, which is a critically endangered species according to the IUCN Red List largely because of its ...
point cat


Silver and golden series


Silver series

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 cold-toned ''
silver tabby A tabby cat, or simply tabby, is any domestic cat (''Felis catus'') with a coat pattern distinguished by an M-shaped marking on its forehead, stripes by its eyes and across its cheeks, along its back, around its legs and tail, and characteris ...
''. On solid cats, it turns the base of the hair pale, making them ''silver smoke''. The term ''cameo'' is commonly used for red silver and cream silver (inhibitor gene (I-O-)) colored coats in cats.


Wide band factors

Silver agouti cats can have a range of phenotypes, from silver tabby, to silver shaded (under half the hair is pigmented, approx. 1/3 of hair length), to tipped silver also called chinchilla or shell (only the very tip of the hair is pigmented, approx. 1/8 of hair length). 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 A polygene is a member of a group of non- epistatic genes that interact additively to influence a phenotypic trait, thus contributing to multiple-gene inheritance (polygenic inheritance, multigenic inheritance, quantitative inheritance), a type ...
trait. File:Poil smoke.svg, Smoke File:Poil shaded.png, Shaded File:Poil tipped.jpg, Tipped (chinchilla / shell) File:Poil cameo shaded.jpg, Cameo shaded (red / cream silver)


Golden series

If a cat has the wide band trait but no silver melanin inhibitor, the band will be golden instead of silver. These cats are known as golden tabbies, or in Siberian cats ''sunshine'' tabbies. The golden color is caused by the '' CORIN'' gene. Shaded golden and tipped golden are also possible, in the same hair length distribution as the silver-gene. However, there is no golden smoke, because the combination of wide band and nonagouti simply produces a solid cat.


Tipped or shaded cats

The genetics involved in producing the ideal tabby, , 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 silver melanin inhibitor gene in some instances does not block pigment, resulting in a grayer undercoat, or in tarnishing (yellowish or rusty fur). The grayer undercoat is considered 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. The genetic influences on tipped or shaded cats are: *
Agouti The agouti (, ) or common agouti is any of several rodent species of the genus ''Dasyprocta''. They are native to Central America, northern and central South America, and the southern Lesser Antilles. Some species have also been introduced else ...
gene. * Tabby pattern genes (such as Ta masking the tabby pattern). * Silver/melanin inhibitor gene ''I/i''. * Golden ''CORIN'' gene. * Factors affecting the number and width of bands of color on each hair, such as the hypothetical wide band gene ''wb''. Resulting in shaded or tipped (chinchilla/shell) pigmentation. * 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). File:RoyalNefertt Serket of AchetAton.jpg, Black silver spotted tabby
Egyptian Mau The Egyptian Mau is a small to medium-sized, short to medium-haired List of cat breeds, cat breed. They are one of the few naturally spotted breeds of domesticated cat. The spots of the Mau occur on only the tips of the hairs of its coat. It is co ...
. File:Cat in greenhouse 2003 (cropped).jpg, Black smoke cat. File:Shaded Cameo Maine Coon male cat.jpg, Cameo (red silver) shaded
Maine Coon The Maine Coon is a large Domestication, domesticated breeds of cats, cat breed. One of the oldest natural breeds in North America, the breed originated in the U.S. state of Maine, where it is the official List of U.S. state mammals, state cat ...
. File:Fat Cat (3365844868).jpg, Black silver tipped (chinchilla)
Persian cat The Persian cat, also known as the Persian Longhair, is a long-haired breed of cat characterised by a round face and short muzzle. The first documented ancestors of Persian cats might have been imported into Italy from Greater Khorasan, Khoras ...
.
File:Murmur's Siberian Cat siberiano golden (cropped).JPG, Sunshine blotched tabby Siberian Cat. File:Goldengirl.jpg, Black golden ticked tabby
British Shorthair The British Shorthair is the Purebred, pedigree version of the traditional British Domestic cat (landrace), domestic cat, with a distinctively stocky body, thick Fur, coat, and broad face. The most familiar colour variant is the "British Blue", wi ...
. File:BRI Golden Garry v. Wahrberg (4488833650) (cropped).jpg, Black golden ticked tabby
British Shorthair The British Shorthair is the Purebred, pedigree version of the traditional British Domestic cat (landrace), domestic cat, with a distinctively stocky body, thick Fur, coat, and broad face. The most familiar colour variant is the "British Blue", wi ...
. File:Cat Golden Chinchilla.jpg, Blue golden tipped (chinchilla)
Persian cat The Persian cat, also known as the Persian Longhair, is a long-haired breed of cat characterised by a round face and short muzzle. The first documented ancestors of Persian cats might have been imported into Italy from Greater Khorasan, Khoras ...
.


Fever coat

Fever coat is an effect known in domestic cats, where a pregnant female cat has a
fever Fever or pyrexia in humans is a symptom of an anti-infection defense mechanism that appears with Human body temperature, body temperature exceeding the normal range caused by an increase in the body's temperature Human body temperature#Fever, s ...
or is stressed, causing her unborn kittens' fur to develop a silver-type color (silver-grey, cream, or reddish) rather than what the kitten's
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...
would normally cause. After birth, the silver fur is replaced naturally by fur colors over the span of a few weeks according to the kitten's genetics.


Fur length and texture


Cat coat hair

Cat fur can be short, long, curly, or hairless. Most cats are short-haired, like their ancestor. The fur can naturally come in three types of hairs;
guard Guard or guards may refer to: Professional occupations * Bodyguard, who protects an individual from personal assault * Crossing guard, who stops traffic so pedestrians can cross the street * Lifeguard, who rescues people from drowning * Prison gu ...
, awn, and
down hair A fur is a soft, thick growth of hair that covers the skin of almost all mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an ...
. The length, density and proportions of these three hairs varies greatly between breeds, and in some cats only one or two types are found. Most oriental breeds only express one single layer of silky coat. However, cats can also have double-layered coats out of two hair types in which the down hairs form the soft, insulating undercoat, and the guard hairs form the protective outer coat. A typical cat coat exists of all three natural hair types, but due to the equal lengths of two of these hairs, the coat is still considered double-layered. Typically, the down hairs comprise the undercoat while the guard and awn hairs make up the basic top coat. Double-coated cats with thick undercoats require daily grooming as these coats are more prone to matting. Double coats are found in for example the
Persian Persian may refer to: * People and things from Iran, historically called ''Persia'' in the English language ** Persians, the majority ethnic group in Iran, not to be conflated with the Iranic peoples ** Persian language, an Iranian language of the ...
,
British Shorthair The British Shorthair is the Purebred, pedigree version of the traditional British Domestic cat (landrace), domestic cat, with a distinctively stocky body, thick Fur, coat, and broad face. The most familiar colour variant is the "British Blue", wi ...
,
Maine Coon The Maine Coon is a large Domestication, domesticated breeds of cats, cat breed. One of the oldest natural breeds in North America, the breed originated in the U.S. state of Maine, where it is the official List of U.S. state mammals, state cat ...
and Norwegian Forest cat. Additionally, there even exist cats which express all three natural types of cat hair in different lengths and structures, which form three different layers. These cats are called triple-coated.
Siberians The Siberians or Siberiaks (, ) are the majority inhabitants of Siberia, as well as the Ethnic group, subgroup or ethnographic group of the Russians. As demonym The demonym ''Siberian'' can be restricted to either the Russian Siberiaks or ...
and Neva Masquerades are known for their unique triple coats, which provides double insulation to withstand their natural cold climate.


Coat mutations

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.


Fur length

Cat fur length is governed by the ''Length'' 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. A rare recessive shorthair gene has been observed in some lines of Persian cat (silvers) where two longhaired parents have produced shorthaired offspring. The ''Length'' gene has been identified as the fibroblast growth factor 5 (FGF5; ) gene. The dominant allele codes for the short coat is seen in most cats. Long coats are coded for by at least four different recessively inherited mutations, the alleles of which have been identified. The most ubiquitous is found in most or all long haired breeds while the remaining three are found only in Ragdolls, Norwegian Forest Cats, and Maine Coons.


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. Some of the rex genes that breeders have selected for are: *
Devon Rex The Devon Rex is a tall-eared, short-haired breed of cat that emerged in England during the late 1950s. The breed is known for its atypical appearance, with an oddly shaped head, large eyes, and the short and wavy coat. History Origin Beryl Co ...
** Mutation in ''
KRT71 KRT71 is a keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horn ...
'' (), the same gene causing hairlessness in Sphynx cats. ''re'' is an allele completely recessive to the wildtype and completely dominant to ''hr'' found in Sphynx. *
Cornish Rex The Cornish Rex is a cat breed, breed of domestic cat. The Cornish Rex only has down hair. Most breeds of cat have three different types of hair in their coats: the outer fur or "guard hairs", a middle layer called the "awn hair"; and the down ha ...
** Mutation in '' LPAR6'' (). A completely recessive allele termed ''r''. * Ural Rex ** Mutation in '' LIPH''. * German Rex ** Provisionally an allele termed ''gr''. Same locus as Cornish, but proposed as a different
allele An allele is a variant of the sequence of nucleotides at a particular location, or Locus (genetics), locus, on a DNA molecule. Alleles can differ at a single position through Single-nucleotide polymorphism, single nucleotide polymorphisms (SNP), ...
. However, most breeders consider the German Rex to have ''r''/''r'' genotype. * Oregon Rex (extinct) ** A hypothetical recessive allele termed ''ro''. * Selkirk Rex ** A dominant allele termed ''Se'', although sometimes described as an incomplete dominant because the three possible allele pairings relate to three different phenotypes: heterozygous cats (''Se/se'') may have a fuller coat that is preferred in the show ring, while homozygous cats (''Se/Se'') may have a tighter curl and less coat volume. (''se/se'' type cats have a normal coat.) This phenomenon may also colloquially be referred to as additive dominance. *
LaPerm The LaPerm is a List of cat breeds, breed of cat. A LaPerm's fur is curly (hence the name "Perm (hairstyle), perm"), with the tightest curls being on the throat and on the base of the ears. LaPerms come in many colors and patterns. LaPerms genera ...
** Provisional completely dominant ''Lp'' allele.


Hairlessness

There are also genes for hairlessness: * h=French hairless cat,
recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...
. * hd=British hairless cat,
recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...
. * Hp=Russian Donskoy and
Peterbald The Peterbald is a hairless cat breed of Russian origin. It was created in St. Petersburg in 1994 from an experimental cross of a Don Sphynx and an Oriental Shorthair. They have an Oriental build with a dominant hair-losing gene. The b ...
, dominant. * hr=
Sphynx cat The Sphynx cat (pronounced , ) also known as the Canadian Sphynx, is a breed of cat known for its lack of fur. Hairlessness in cats is a naturally occurring genetic mutation, and the Sphynx was developed through selective breeding of these anim ...
,
recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...
. Identified on
KRT71 KRT71 is a keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horn ...
. 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 Guard or guards may refer to: Professional occupations * Bodyguard, who protects an individual from personal assault * Crossing guard, who stops traffic so pedestrians can cross the street * Lifeguard, who rescues people from drowning * Prison gu ...
, awn, and
down hair A fur is a soft, thick growth of hair that covers the skin of almost all mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an ...
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 Ornithine transcarbamylase deficiency also known as OTC deficiency is the most common urea cycle disorder in humans. Ornithine transcarbamylase, the defective enzyme in this disorder, is the final enzyme in the proximal portion of the urea cycle, r ...
in mice.


Loci for coat colour, type and length


See also


References


Further reading

* * *


External links


Cat Genetics articles in plain English
{{DEFAULTSORT:Cat Coat Genetics Genetics of cat coats Cat genetics Mutation Animal hair