BLUE-EYED BREEDS / DOMINANT BLUE EYE (DBE)
This page describes blue-eyed cats in colours other than white, bicolour or colourpoint (Siamese pattern). Some of the Russian breeds have variable spelling when the breedname or cat name is translated into the Roman alphabet. This information will be updated as more of the gene's effects become apparent. There is currently no DNA test for the gene(s) involved. Firstly, there is no single "DBE" mutation so it is important to know which gene you are discussing/working with as they should not be crossed to each other (until the genes are identified). So you need to distinguish "Altai BBE," "Barnaul DBE," "Australian DBE" or "unknown DBE" etc. Secondly, the modern blue-eye breeds are not Ojos Azules - that breed is extinct, it never made it out of America and the more recent mutations do not have the same side-effects.
Blue-eyed non-white cats were discovered almost 20 years apart and on almost diametrically opposite regionss of the globe: the California Ojos Azules and the Kazakhstan Altay Blue-Eyes. In both cases, outcrossing showed that the blue-eyed genes were dominant type over the ordinary yellow, copper or green colours. In both cases there is at least a tiny white marking, most often at the tail-tip, but it might also be a white locket, white back paws or a white smudge on the face. Due to incomplete penetrance of the gene, the size of the white marking ranges from just a few hairs to a large white patch (in homozygotes).
It is currently assumed - though DNA analysis is still ongoing - that the Altai blue-eyes gene is an allele of the white spotting gene and that it is “weakest” in comparison to the usual white spotting gene which causes the familiar bicolour range and it may also interact with the white spotting gene. Genes encode for proteins and any change to a protein can have other effects.
ALTAI / ALTAY
Olga Volynska was the first to notice blue-eyed cats at a factory, but her report was largely ignored because conventional wisdom said blue-eyed colour cats could not exist. A couple of years later the Ojos Azules was reported in a TICA journal. Lyubov Borisovna Zikeeva from Ust-Kamenogorsk noticed the blue-eyed cats on the street, and decided to find out more. The first Ust-Kamenogorsk Altai was an ordinary domestic semi-long-haired cat, with white marks, with the right eye blue, and the left eye yellow-green. If the Ojos Azules could be recognised as a new breed on the basis of eye colour and small white marked, then the Altay could be recognised too! The cat magazine "Friend" (issue No. 8/9 August-September 2004) reported that the first owner of an Altai cat was Vera Sokolova, who lived in the city of Ust-Kamenogorsk in Kazakhstan. In 1995 she owned a black cat called Fyodor who had blue eyes and a white tail-tip, a phenotype that resembled the American Ojos Azules discovered in California.
Lyubov Borisovna Zikeeva began talking with Fyodor's owner, and from him she embarked on the creation of the Altay blue-eyed breed which has lasted more than twenty years and continues today. After many unsuccessful matings, Zikeeva L.B. was finally rewarded by the birth of a blue-eyed cat named Ariel, which is considered the progenitor of all living Altai cats. After breeding a blue-eyed kitten, Zikeeva mated blue-eyed to blue-eyed to get a wholly blue-eyed litter. This is when she discovered that homozygotes were deaf. Kittens with one eye blue and the other green-yellow are heterozygotes (the gene is dominant with incomplete penetrance) and from the point of view of deafness should be used in breeding, while homozygotes with two blue eyes should not. Line-breeding for generations is used to fix the traits, but in the third generation an outcross to green-eyed European Shorthair is needed to prevent inbreeding depression.
Early Altai cats bred by Zikeeva L.B. include Roxy/Roksi (tabby with white hind feet and white throat); Grand Yegan (tabby, white cheeks and chin); Ekatarina (tabby with white hind feet and white throat); Nils (tabby, white paws, muzzle and throat). A blue-eyed Altai bred by Lubov Borisovna is Ellie Darling (black tuxedo pattern). By 2016, there were still not a lot of breeders working with the Altai. One of them is an experienced breeder and veterinarian Elena Lobasova (Murmansk, Russian Federation), the other is Anna Kalinichenko (Zaporozhye, Ukraine). “Wonderful Aquamarine” cattery, (Grin L.V. and Kalinichenko A.O., Ukraine), began breeding Altay (Altai) cats in 2016 under the framework of the WCF registering body. Wonderful Aquamarine studs in 2017 were Efimenty Veles Darlin (silver spotted tabby with blue and green odd eyes) and Sonny Sweet Dreams of Wonderful Aquamarine (blue-eyed red). A queen shown on the website is Missy Nikital (blue-eyed classic tabby tuxedo pattern).
It is undesirable to cross two blue-eyed or odd-eyed Altay cats as this can lead to deaf cats with a high degree of white in the coat. Blue-eyed Altay should be bred to non-blue-eyed Altay variants, preferably to green-eyed variants. This produces strong and healthy offspring and a mix of blue-eyed, odd-eyed and green-eyed offspring. Unlike the American Ojos Azules, Altay cats do not have gross physical deformations associated with the mutation. Along with the Ukrainian Topaz breed they are mistakenly called “Russian Ojos Azules” by English speakers. Altay Blue-Eyed were used early on with Topaz which combines two different lines of blue-eyed cats, but which differs in white marking pattern and sometimes having black-roan fur.
In the Altai, the bright blue eyes, or odd eyes, can be combined with non-white cats. The breed code is ALT and it is recorded in the register of new breeds and cat colours by the Tribunal Commission of the SFF in 1999. In that year, a draft standard was created and registered for the Altai Blue-Eyed (also known as ABE for short). This old standard called for a rounded wedge shaped head, cheeks not very pronounced, straight nose, the transition to the forehead being smooth without indentation (the 2018 standard calls for a distinct stop), strong jaws and rounded chin. Eyes are round and medium sized, set slightly obliquely and are blue or odd-eyed. Ears are small with slightly rounded tips, slightly taller than they are wide at the base (the 2018 standard calls for large ears, very wide at the base). Body is rectangular, medium sized, strong, flexible and muscular. Legs are medium length, and paws are round. The colours are solids, tabbies or torties with or without white, and there is a distinctive white tail-tip which may extend to one-third the length of the tail. Blue-eyed Altai without any white are the most prized. Fur is short, dense, glossy and springy with a well-developed undercoat.
The experimental breed has been exhibited in non-competition classes or as household pets. Breeders considered this unfair because some Altai pedigrees now have 5 or 6 generations of Altai-to-Altai breeding and are maintained and registered, whereas household pets can be mongrels. A WCF standard for the Altai was drawn up in 2018 (Anna Kalinichenko) to promote the Altai as a distinct breed and differentiate it from the Ukrainian Topaz and Celestial British. It calls for a medium to large, muscular and sturdy cat with a dense coat, vivid blue eyes and non-pointed colours. Its vivid blue eyes are wide-set, oval and slightly tilted - their outer corners being above the level of the outer base of the ears. The eye-colour should be rich, vivid and uniform blue, blue-green or green, or odd-eyed with good contrast between the two eye colours. Any eye colour is permitted, but blue and green colours are preferred for exhibition-quality cats. Strabismus (crossed eyes) is a disqualifying fault.
The head is round (a modified wedge with the width being almost equal to the length), the muzzle is fairly short, the chin is firm, the whisker-pads and cheekbones are pronounced. The ears are large, very wide at the bases and low-set with tufted tips and ear furnishings. The forehead is rounded and the profile has a distinct stop. The neck is long, strong and muscular and proportionate to the head and body. The body is medium length, strongly boned and compact with a broad chest. Males are larger than females. Limbs are strong, medium-long and slender (not cobby) with the hind-legs being slightly longer than the forelegs. The paws are oval; polydactyly is not permitted. The tail is medium length (long tails are also permitted, but not short, kinked or bobbed tails) and well-furred. The fur is especially thick on the back, legs, muzzle, cheeks and ears, sides, and tail, and there must be no signs of baldness. The coat can be short or semi-long. All colours are allowed, including bicolour, although cats with as little white as possible are preferred. Solid white and colourpoint are not permitted.
Until 2030 it will be allowable to out-cross Altai cats with American and European Shorthair, British Shorthair, Bengal, Abyssinian, Somali, and Siberian to produce the correct conformation, but after May 1, 2030, out-crossing will be prohibited and only Altai-to-Altai matings will be allowed. However, breeding together two blue-eyed or odd-eyed Altai cats is not permitted because it may lead to deafness and white coats in cats homozygous for this dominant gene. Regarding deafness, the Altai gene has similar risks to “blue-eyed white” and “blue-eyed high-white bicolour” because the absence of melanocytes affects inner ear development as well as eye colour. Blue/odd-eyed Altai may only be bred to non-blue-eyed Altai (preferably green-eyed) which will produce litters with a mix of blue-eyes, odd-eyes and green-eyes.
Altai cats are friendly and adapt well to their environment. They are active, energetic, sociable and affectionate and take an interest in everything going on around them. They do not require excessive space and they get on well with other animals. No special care or attention is required beyond the normal care of a short-haired cat.
BARNAUL AND OTHER BLUE EYED MUTATIONS FROM EASTERN EUROPE
Test matings of another blue-eyed cat found in the Altai region proved to be a different gene and have been investigated by Rano Makarenko. Why are different blue-eyed genes suddenly appearing in Russia and Ukraine? Most likely they have always been in the random cat population, but no-one took any notice. With Eastern European countries developing their own home-grown cat fancies and developing local cats into recognised breeds, these blue-eyed cats are now getting the attention they deserve.
Rano Makarenko also found several blue-eyed cats from Aktau, a city on the Caspian sea and there are photos of others from cat rescues in the same region.
Elena Paukova is breeding blue-eyed Munchkins since 2019.
Elizaveta Lipovenko has a spontaneous mutation in her British Shorthairs, producing blue-eyed and odd-eyed chinchilla/silver shaded and tipped golden cats.
The Topaz is being developed by Iryna Merzlenko in Nikital cattery in Ukraine starting in 2016. It is the Asian equivalent of the Ojos Azules and is also mistakenly known as the Russian Ojos Azules. It is not related to the American Ojos Azules. The first cat came from Kazakhstan (Eastern Altai) and is the Altai Goluboglazaya breed (Altai Blue Eyes - dominant gene with incomplete penetrance i.e. some cats can have the gene but not display it). The second cat came from Russia, 3,000 km from the first cat and was dubbed the Russian Ojos Azules (Russian Blue Eyes) because it resembled the American breed (which was never exported from the USA). Whether it is a mutation of the same gene as the unrelated American cats can only be found out by genetic testing. Unlike the American breed, the Russian cats do not appear to have gross physical deformities associated with the mutation, however most homozygotes are deaf.
The preliminary standard for the Topaz calls for a small or medium-size, compact cat. They are not high on paws. The tail is straight, round and thick. The head is medium sized and rounded, with a well-developed chin and good jaw (no more than 2mm undershot). The ears are small or medium size, broad at the base; the tips may have a small brush. The Shorthair Topaz has a short, glossy, close-lying coat and almost no undercoat. The Longhair Topaz has a long outer coat and a small amount of undercoat, a very long and fluffy tail, fluffy britches and a small neck frill. To prevent mixing with blue-eyed whites, the coat colour can be anything except pure white or predominantly white. If there is more than 25% white, the tail tip should be a solid dark color.
The eyes are large or very large, round, wide open and expressive. The eye colour is the distinguishing feature. Eyes must be blue – either both eyes or odd eyes. The blue ranges from deep, bright blue through to violet blue. Cats with red pupils may be used for breeding. In odd-eye Topaz, the non-blue eye is often a shade of green or yellow-green.
In later generations, black-eyed Topazes occurred. These had irides so dark-coloured that they are black. On close inspection, they appear to be a very dark olive green. Around 6 kittens with black eyes were born several times from certain combinations of parents, but none of them survived, possibly due to weak immune systems. The black eye colour was an indication of an undesirable combination of genes so Iryna sterilized the lines that produced this trait.
The Topaz is a playful, affectionate, people-centred cat.
Acceptable outcrosses are Bombay (Asian Shorthair), British Shorthair, European Shorthair, Altai Goluboglazaya. Crossing to the Scottish Fold is not permitted. It is not advisable to mate two blue-eyed cats together in case this gene has similar side-effects to the gene found in the unrelated American Ojos Azules. Topazes have been exported to American breeders, the Gobbles, who intended to register them as Ojos Azules even though the gene is different. Having obtained the Topaz cats for breeding in the USA, contact with the breeder was blocked. Cats have also been exported to France (Nikital line) and Germany to found new varieties there.
According to Iryna, the percentage of deformities and stillbirths is about the same as in any other breeds of cats. In the case of homozygotes which have a high degree of white, the only drawback is deafness, but even in this case not all high-white Topaz cats have this. Iryna's female Christina, which has the same "white shirt and dark trousers" pattern (dubbed Panda pattern) as Panda, has a normal hearing while Panda does not. A more extreme pattern is "chipmunk" - all white with a short colour stripe along the backbone. These "high white" patterns occur only in homozygotes and the pattern is random, not inherited consistently. Homozygotes are not used in Topaz breeding and the breeding combinations are selected in order to avoid them. The other anomaly is that homozygotes tend to have a shorter nose and sometimes deformation of the nasal mirror (deviation of the nasal septum), but this does not affect breathing or lifestyle. Because the Topaz cats are derived from two distinct blue-eyed populations that occurred 3,000 km apart, there may well be two different genes at play. So far, only one breeder who crossed a Russian blue-eyed cat with another breed in order to introduce the blue-eyed trait, found higher than expected mortality in kittens, but this might be due to factors unrelated to the eye colour gene.
One of the blue-eye genes involved also affects coat colour. The colour-and-white-blanket patten (Panda pattern) which looks like a cat wearing a white shirt and coloured trousers is linked to one of the blue eyed genes and to deafness in some of the cats. These cats can also change from black to silver grey or black-roan in a manner resembling the karpati pattern. Karpati, roan and rigntail are all desirable characteristics in the developing breed.
AMERICAN OJOS AZULES (EXTINCT)
Cats with vivid blue eyes were discovered in New Mexico among feral cat populations in the 1980s. In 1984, a blue-eyed tortoiseshell named “Cornflower” was bred to non-blue-eyed males resulting in litters of blue-eyed kittens. This showed the trait to be dominant. The breed was named “Ojos Azules,” this being Spanish for 'Blue Eyes'.
Foundation female “Cornflower” (tortoiseshell) was found in an animal shelter in New Mexico by Annie Gass who brought her to the attention of TICA’S chief geneticist, Dr Solveig Pflueger. Cornflower had been found in a feral colony and had brilliant blue eyes. When bred to unrelated males, the kittens inherited this eye colour. During the 1990s, breeders worked to develop the Ojos Azules breed. TICA recognized the Ojos Azules as a breed in 1991 but it remained a rare breed with only 10 known individuals in 1992. The gene that caused the blue-eyed trait was found to be more complicated than a straightforward autosomal dominant. Breeding was slow because of small litters and the risk of deformities.
All of Cornflower’s descendants had a distinct appearance: wide-set eyes, distinct cheekbones and plump whisker pads. They often had white tail-tips regardless of their coat colour. Several Ojos Azules were odd-eyed, which was accepted in the standard to ensure diversity and because homozygous kittens had cranial deformities, white fur, a small curled tail and they died in the womb. The cranial deformities except they were described as similar to those seen in “double merle” dogs and this led Solveig to try to investigate the gene without breeding any further kittens. When the gene is heterozygous, these lethal genetic mutations do not occur. This means it is undesirable to mate two blue-eyed Ojos Azules together. The deformities can be avoided by breeding a blue-eyed cat with a non-blue-eyed cat. This produces a litter with approximately 50% blue-eyed kittens and 50% non-blue-eyed kittens.
For comparison, merle in dogs is an autosomal, incomplete dominant mutation causing mosaic fur and blue or odd eyes. In dogs it is a mutation in the region of CFA10 that contains the SILV(aka PMEL) gene. Merle dogs (heterozygotes and homozygotes) can show a wide range of hearing and visual abnormalities similar to human Waardenburg syndrome. To complicate matters, the merle gene can vary in size (repetitive DNA), with the pattern depending on the size of the merle gene, and different cells can have different size merle genes. Double merle in dogs causes defects such as deafness and blindness; deformed, tiny or absent eyeballs; 'starburst' pupil, where the pupil appears to have spiky projections and there is light-sensitivity. They have a high degree of white, hence the gene is also called “lethal white.” The stillborn homozygous Ojos Azules kittens also had white fur and one heterozygous male had light sensitivity and problems contracting the pupils.
Cornflower’s daughter “Los Nuevos Iris Polychrome” (Polly) had eyes that were sapphire blue, patched and flecked with gold. Polly’s daughter “Margaree’s Billie” (Billie) was a red mackerel tabby with deep amber eyes but she had the facial features of an Ojos Azules. When bred to a non-Ojos Azules cat she produced four kittens, three blue-eyed and one odd-eyed, proving the theory that some genetically blue/odd-eyed cats did not exhibit the blue eyes. In the more recent blue-eyed Russian cats this is known as “latent” blue eyes (equivalent to “”cryptic merle” in dogs). Although a dominant gene cannot be “carried” it seems that the dominant trait is not always expressed (perhaps because these are all “white spotting” genes and the white spots need to be around the eyes in order to get blue eyes).
Orange eyed Billie was a non-standard Ojos Azules and could not be exhibited, but she was useful to the breeding program because she was genetically blue-eyed. Solveig kept Billie’s kittens, but it was around this time that breeders became aware of the problem of breeding Ojos Azules cats to each other, regardless of their eye colour. The limited breeding data found that 9.5% of non-standard Ojos Azules (orange eyed “latents”) also had birth defects. Some breeders left the breeding programme at that point because there was an unacceptably high risk of producing deformed kittens and because of the number of non-blue-eyed cats that would be produced when avoiding homozygous kittens.
Breeders could avoid the lethal defects by only breeding an Ojos Azules cat to cats of other breeds, for example a non-pedigree Domestic Shorthair with a similar facial type (because an Ojos Azules is more than just a blue-eyed cat). The resulting outcross litter should be 50% blue eyed cats and 50% not blue eyed. This was commercially and ethically non-viable because around half the kittens would need to find pet homes every time and pet-buyers would want the novel blue-eyed cats. The distinctive facial conformation was likely to be lost over just a few generations. Less scrupulous breeders would ignore a safe breeding methodology and would breed blue/odd-eyed cats together to ensure a high percentage of blue-eyed offspring, many of which would have physical deformities.
The Ojos Azules was an incredibly beautiful and friendly cats. A single longhaired red and white male, “Jeepers Creepers,” was fearful and timid, but he had a problem with contracting pupils and may have been visually impaired. Visual impairment was not found in any other Ojos Azules. While blue-eyed cats turn up at random in the USA, these are unrelated to Cornflower’s line and are not Ojos Azules breed – the Ojos Azules was also characterised by its conformation.
Unlike the blue eyes seen in white cats, some bicolours and colourpointed cats, the Ojos Azules gene was not linked to a particular fur colour or pattern. It also produced a deeper shade of blue than in colourpointed cats. It was also not associated with squinting or crossed eyes (seen in some colourpointed cats) or with deafness (seen in some blue-eyed white cats). Breeders saw this as an opportunity to breed blue-eyed tabbies, blue-eyed solid black cats etc. One indicator of the presence of the Ojos Azules gene was a flattened tail-tip. Unfortunately, the gene was also linked to birth defects; kittens inheriting two copies of the gene being deformed or stillborn. It wasn’t clear if the deformities were caused by the Ojos Azules mutation itself or by an adjacent or linked gene. Solveig Pflueger investigated these issues and small-scale breeding then resumed in the hope of breeding Ojos Azules without the lethal genetic defects.
The blue colour is described as having depth (hue) greater than that seen in a Siamese. Regardless of the causal mutation, blue eye colour in cats is due to the absence of melanin in the iris The depth of colour is due to polygenetic variations and many show cats (Siamese, blue-eyed whites) have equally deep blue eyes. The contrast of blue eyes against a dark or patterend coat is the main attraction.
The breed was accepted for registration by TICA in 1991 in both long-haired and short-haired varieties. By 1992, only ten Ojos Azules were registered. The breed standard was last updated in 2004. Only those cats with deep blue eyes are termed the Ojos Azules, the other cats in the breeding programme are “variants” and are important for the health of the breed. The standard says “Eye colour should be clear blue or blue-grey, not turquoise or lavender. In odd-eyes, the second eye may be gold, copper, or green. A deeper eye color is preferred.
The Ojos Azules standrd called for a medium-sized cat with large, round blue eyes. The neck is arched. The tail is proportionate to the cat's body. The head is somewhat triangular in shape with a slightly rounded forehead, an angular muzzle and a slight nose break. The coat is short, fine, soft, silky, and shiny without too much undercoat. All colours are allowed and white markings on the extremities (tail-tip, hind paws, and muzzle) are common, as is a sprinkling of white hairs. Because the eye colour is due to lack of melanin pigment, white markings elsewhere may be inevitable. Particolours must have a white tail tip as this is an indicator of the Ojos Azules gene, however a white locket on the chest or white spot on the belly are considered faults. Solid white is not desirable because it is visually the same as existing blue-eyed white cats. The cats bred were active, friendly, and affectionate.
BLUE-EYED GERMAN LONGHAIR
Nicole Ehrenboth (Cattery vom Leineufer), Hannover, Germany breeds the German Longhair (DLH – Deutsche Langhaar) and in collaboration with Miriam Volk and Anette Lanetzki (Cattery von Barnstorf) has introduced the Altai blue-eyed gene into the German Longhair. The breeding programme for blue-eyed Deutsche Langhaar began in 2017 using Muscari of Aerlin, a Celestial British.
CELESTIAL BRITISH (BLUE-EYED BRITISH LONGHAIR)
In August 2016, the LOOF (French registry) Committee of Standards approved the introduction of the blue eye colour in the British Shorthair by Chatterie d’Aerlin (Sophie Guittonneau) and Chatterie de la Grace (Stephanie Jimenez) (Chatterie de la Grace left the Celestial breeding programme later on). In 2017, the partipating breeders included Lazycat Compagni (Vanessa Marmolejo), Vivarnon Cattery (Ghislaine Michel-Paulsen) and Chatterie les Gummis en Sucre (Sylvie Huber). The blue-eyed cats used were considered European Shorthairs and the progeny of the matings to British Shorthair/Longhair cats were registered as XSLH or XLH (experimental SH/LH). All the blue eyed kittens born at Aerlin cattery are tested with BAER for hearing because deafness has been observed in some of the homozygotes. No deaf cats are used in further breeding.
In October 2015, the black smoke Altai “Lagoon” died of pneumonia before joining Chatterie d’Aerlin from Ukraine so her black smoke sister, “Kaena” was sent instead. Kaena was not blue-eyed (light green eyes) but carried the incompletely dominant gene and would produce blue-eyed kittens when mated to “Lazuli” from the same breeder in Ukraine. Kaena expressed the Altai blue-eye gene in a minimal way - a white locket, white belly spot, some white whiskers and the light green eyes. Kaena’s sire had blue eyes. Lazuli was a brown mackerel tabby, with one green eye and one blue eye, from a blue-eyed sire. He carries longhair and is negative for the white spotting gene.
January 2016 - Lazuli x Igritte (cinnamon tortie BSH) produced 6 kitten, two of which expressed the Altai gene: Mirage of Aerlin, blue-eyed red self male (genetically mackerel tabby) and Muscari odd-eyed red mackerel tabby long-haired male, longhaired. The other 4 kittens were red males, “Mermite” and Mogwai” (longhaired), “Maya” - black tortie female and “Malice,” long-haired brown tabby female.
18 July 2016: Kaena x Lazuli produced 7 kittens including 2 homozygotes: “Matcha” - black longhair males, not blue-eyed. “Merlin” – black longhair with two blue eyes and white spots on the nose, back legs and tip of the tail. “Mavi” - A female black smoke longhair with a large white marking on the face and two blue eyes. “Mistral” - silver tabby male with white spots on the forehead, legs and tip of the tail, and blue eyes. “Maïdi “ - female brown tabby without white or blue eyes. “Miyuki” - a solid white longhair with a few coloured hairs in the lower back, and blue eyes (homozygote, deaf, not bred from). “Mochi” - bicolour mostly white, with a big colored spot on the lower body and blue eyes (homozygote, not deaf) – this is the pattern known as “panda pattern” in the Topaz breed that was developed from Altai cats.
In August 2016 Ilvy Freyja of Preston (copper-eyed black British Longhair female b. October 2010) x Lazuli produced 4 kittens. Matahina of Aerlin (female) was solid black with a small white locket and odd eyes, but sadly died. Maïdee (female) was a black longhair with a white marking on the upper lip and blue eyes - she has gone to David Karamatic in Australia to introduce blue eyes into his solid colour Munchkins. Mauve (female) was a black female shorthair with white lips and chin, and two blue eyes. Monky was a brown spotted tabby shorthair male with non-blue-eyes. Mauve d’Aerlin went to Chatterie de la Grace to continue the breeding programme.
In April 2017 Lamu (Tortie BSH) x Muscari produced 3 red kittens, 1 black and 1 chocolate torbie - 4 of which had the coveted blue eyes. Also in 2017, Ilvy x Muscari produced 2 blue-eyed kittens and one non-blue-eyed. The charts below are from informatin kindly provided by the breeders.
In 2019, a new Celestial line began with Azure Dream Jimmy (red odd-eyed shorthair Altai gene cat), Jimmy is descended from a female BRI x male Altai gene cat from Kazakhstan).
The "Heavenly Eyes" programme uses one of Seymour's descendants, Nikital Daisy, from Iryna Merzlenko, as its foundation.
BRITISH ICE SHORTHAIR/LONGHAIR
These are basically British Shorthair/Longhair cats with dominant blue eyes, odd eyes or sectoral heterchromia (iris split into different colours). Solid white is not accepted. It is only recognised by the Rare and Exotic Feline Registry and apart from the addition of sectoral heterochromia appears equivalent to the Celestial. It can be crossed to the British, non-folded variants of Scottish Fold (Scottish Straight/Scottish Shorthair) and straight-haired variants of Selkirk Rex (Selkirk Straight).
BLUE EYED PERSIANS
Another breeder working with the blue-eye gene is Chloe Alaska (Bordeaux, France) who has Nikital Gioia from Ukraine (bred by Iryna Merzlenko). Mated to Ylang Ylang, Chloe’s black Persian stud, Gioia’s first litter comprised 7 kittens, but a black silver tabby female with a node on her tail was stillborn. The other 6 were perfectly healthy and included a blue-eyed kitten and 2 odd-eyed kittens. 2 kittens in the litter had a tiny defect on the last vertebra of the tail, but Chloe had previously seen this in unrelated non-blue-eyed) cats. The blue-eyed cats, included Gioia, have had hearing tests and have perfect hearing. Niagara has a tiny white spot under his nose (right side).
BLUE-EYED RAGAMUFFIN (BLUEBERRY MUFFIN)
In 2017, Josee Rodrigue of Gatineau, Quebec, Canada, imported Alaska's Ninja of RagaMania, an odd eyed male from Chloe Alaska (who had Nikital Gioia bred by Iryna Merzlenko), to introduce the Altai blue eyed gene into theRagaMuffin. Josee breeds the authentic RagaMuffin, and her cats have more than 5 generations of RagaMuffin x Ragamuffin breeding. Ninja is all Black with a white spot on his chin and couple of white hairs on his neck. One the first offspring was black with a white locket on his chin and his eyes are different shades of gold that both reflect red. This shows that he has the dominant blue-eye gene, but it has not shown up in the eye colour.
This division of Josee’s RagaMuffins are called Blueberry Muffins. The offsprings are registered as experimental with the CCC in Canada as experimental. Three litters were born in 2019, some having blue eyes.
There are also blue-eyed sepia and blue-eyed mink Ragdolls variants where the blue eye colour appears to have come from blue-eyed Burmese ancestors.
BLUE EYED MUNCHKINS
Blue eyes have been introduced into the Munchkin breed (self colours only) in Australia by David Karamatic. He imported Celestial British "Maidee" from Sophie Guittonneau and is the first of that gene in Australia.
BLUE-EYE ELF & BLUE-EYE SPHYNX
Jacqui Stevens (Sphynxwillow Sphynx Elf & Bambino Australia) imported Violaflower's Vanessa (born 11-April-2019) in 2019. Vanessa was bred by Tatyana Lackies in Germany and has the dominant blue-eye (DBE) gene. She is a fully registered Elf in Australia with ANCATS and will be on the show bench as soon as shows resume in Australia after the Covid-19 pandemic shutdown. Vanessa is part of Jacqui’s Elf breeding program in Australia. Jacqui is the foundation breeder of the Elf in Australia and has worked to get them fully recognised and registered to Championship Status. Vanessa is an important part of that program. Jacqui will also be importing a DBE Sphynx later in 2021 from Russia from a breeder that is past the 5th generation of that variety. These are the first DBE in a hairless breed in Australia. Vanessa is a blue tabby & white harlequin-pattern with blue eyes and DNA tests show she is negative for the colourpoint blue eye gene.
Vanessa had a litter early in 2021 and gave birth to one DBE female kitten named Sphynxwillow Azure Ono, a blue tabby and white, which has gone to Jane Cockerton (Born Bold), a registered breeder, as part of the breeding programme.
The idea of creating a unique cat resembling a panther (leopard) with blue or odd eyes, came from one of the founders of the Altai breed, Anna Kalinichenko, in 2016. The foundation cats were an Altai blue-eyed cat called Sapphire and two melanistic Bengal cats called Agatha and Bager. Sapphire and Agatha produced the first Pantherina, named Lynx Ellie. In 2020, the preliminary standard for the Pantherina breed was created and in 2021 this experimental breed began to gain popularity.
The Pantherina gives the impression of a graceful wild cat with spotted, rosetted or melanistic coat, with any white restricted to tail-tip or feet. The large, wide-set, oval eyes should be deep blue or odd-eyed (one deep blue, the other emerald or amber) due to the altai dominant blue-eyes genes. Size is medium to very large, imposing in appearance, body is well-balanced, strong, muscular and flexible and the demeanour is active, energetic and alert. The head is small in relation to tye body and is a modified wedge with smooth outlines, longer than it is wide. High cheekbones, wide muzzle with prominent whisker pads, slight curve between forehead and nose, just above eye-level. Ears are medium size, broad at the base, directed forward and rounded at the tips. Neck is muscular, long and appears thick in relation to the head size. The body is not as elongated as an Oriental. Legs are medium length and muscular with large, round paws; hind legs are somewhat longer. The tail is rather long, thick at the base and tapes to a rounded tip. The coat is short, dense and silky with a glittering effect. The spotted pattern is random or horizontally aligned, preferably with horizontal shoulder markings. The marble pattern should also be horizontally aligned, with a darker centre of the markings compared to the background colour. In both cases there should be spotted markings on the belly. Striped pattern also occurs. Colours are brown (warm-toned/tawny background with black, brown or chocolate markings), silver with highly contrasting pattern (dorsal region may be a little darker than sides), charcoal (cold-toned brown with darker face and pale “glasses” around the eyes), blue (metallic blue-grey with dark blue-grey pattern) or melanistic (solid black with ghost markings visible in certain light). Because the blue eyes are associated with a white spotting gene, blue-eyed homozygotes may have any amount of white present (except solid white), and odd-eye cats may have white feet or white boots. A white tail-tip is desirable in odd-eye cats and white spots on the belly and muzzle are acceptable. Faults include short, bobbed or kinked tail, polydactyly. Cats with 2 blue eyes (homozygotes) may not be bred together. Until 2050, outcrossing to Altai, Bengal and Maine Coon is allowed to maintain or improve conformation and gene pool.
AUSTRALIAN DOMINANT BLUE-EYE MUTATION
David Karamatic is working with a dominant blue eyed mutation discovered in tropical north Queensland, Australia. He was notified of some “free to good home” kittens that had retained their blue eyes. There were four in the litter three with blue eyes and one odd eyed. At first he thought it was the influence of white-spotting, but the odd eyed kitten was nearly completely black. He obtained the last kitten available, and at seven months old its eye colour is very deep and vivid almost violet. It would be interesting to see if this gene is the same as the Celestial / Topaz or perhaps on the same locus. David first wants to find out if the mutation is the same as his imported French Celestial (Maidee, from Chatterie d’Aerlin), and he is also submitting samples for DNA tests.
BLUE-EYED SILVER AND GOLDEN BRITISH SHORTHAIRS
Two golden British Shorthairs with blue eyes - Ermine Trace Athens and Ermine Trace Zarafina) both trace their ancestry via Ermine Trace Nadeya to a very blue-eyed chinchilla point male, Shoenwig Teodoro and his mother Schoenwig Mona Lisa, also a very blue-eyed chinchilla point cat. Mona Lisa’s mother is Passimilla’s Upper Class, also a very blue-eyed chinchilla point cat. It is possible that in selecting for very blue eyes, a gene mutation may have been overlooked. Because pointed cats have blue eyes, it is also possible the mutation could have been hiding in plain sight.
Information on another dominant blue-eyed mutation, with no apparent links to the Ermine Trace cats, has been researched and provided by Sarah May. It occurred in the Nanotigr cattery. When the breeder gave up breeding, the blue-eyed cats went to Angela Savenko (Hermes Cattery), a breeder of bicolours, and were also bred to Selkirk Rex. She sold a blue-eyed black-tipped British Shorthair to British SH/Selkirk Rex Cattery annealtindan (Kaan Altindan) in Turkey. It isn’t known whether the Capuchino line descended from Roddiffer have produced any cats with odd eyes. The breeder believed them to be “latents” (having the blue-eyed mutation, but not displaying it), but they had no white lockets which are normally indicators of the mutation.
BLUE-EYED BURMILLAS, BURMESE AND TONKINESE
Blue eyed kittens sometimes occur in Burmilla (derived from Burmese x Chinchilla ancestors) lines with browns and chocolates in the pedigree. Although they don’t match the breed standard (green eyes), they were usually the first kittens to be sold. The blue eyes likely trace to a Himalayan ancestor many generations ago. DNA tests have identified the colourpoint (Siamese) gene in some Burmillas. In the 1980s, Himalayans/Colourpoint Persians and Chinchillas were crossed to produce the lynx and smoke point series Himalayans/Colourpoint Persians. Genetically chocolate cats (and chocolate carriers) were used in the breeding programmes and some Chocolate Chinchillas/Goldens were registered without stating they were chocolate. Aquamarine eyes have also been seen in Burmillas, where the Burmese sepia gene meets up with the colourpoint gene. Because many breeders breed for light coat colour without markings, aquamarine eyes (linked to mink colour) are less important than clear silver. This also means that Burmillas have become lighter in colour than the original British cats. DNA testing on some Burmilla lines also identified the white gloving gene is present although it is not visibly expressed.
There is a line of blue-eyed Tonkinese from Joan Bernstein who bred some blue eyed sepia variants. Descendants of her cats continue to produce blue eyed sepias and blue-eyed minks.
CHENGGE MAO CATS, CHINESE DOMINANT BLUE-EYE MUTATIONS
In November 2020, Rano Makarenko sent me information about some Chinese cats known as Chengge mao. The variety lives in Chengge and the breeder selects them for mouse-catching and is not at all interested in genetics questions. Originally, the Chengge mao was white with a coloured tail, symbolic of good mood and richness because of a resemblance to a rice bowl with chopsticks (Van pattern without head markings). Then the cats started producing blue eyed cats from the white cats with coloured tails.
Blue-eyed solid colour cats (often red-colour cats) are known to be bred and sold in part of China, but there is no control over the breeding and the gene(s) involved are unidentified. One breeder reported a hgh mortality rate with 50% of kittens from his blue-eyed red ticked female mated to a blue-eyed red classic tabby male being solid white and stillborn/dying at birth. Though there could be various reasons for this, there is a possibility of 2 incompatible blue-eye genes being bred together. Breeding the blue-eyed red-ticked female to an unrelated normal-eyed cat and then breeding a male offspring back to the red-ticked female would produce kittens with only one blue eyed mutation (and could also show if the stillbirths were due to a reproductive issue with the female herself.)
This is a blue-eyed breed being developed in Australia by crossing the bobtailed, curl-eared Highland Lynx (TICA name "Highlander") and Altai Maine Coon (a Australian developmental form of European-style Maine Coon with dominant blue eyes). It is a strongly-built, muscular cat with large, loosely curled ears with lynx-tufts. The loose curl of the ears is found in cats heterozygous for the curl gene. The paws may be polydactyl and the tail may be long, or any length of bobtail. It has a broad, strong muzzle and wide-set oval eyes. Both longhair and shorthair forms are bred. Only the black-based colours are accepted and may be solid, tabby or silver/smoke. White spotting is undesirable, but may be unavoidable. In order to maintain the desired conformation, it can only be outcrossed to the Maine Coon (European style).
The photos have been provided by Jake Macmillan and the breed is currently (2021) registered with REFR. The Altai Maine Coon is accepted in Australi as a developmental breed.
OTHER BLUE-EYED CATS & HISTORICAL REPORTS
The blue-eyed trait turns up surprisingly often in random-breeding cats. A grey bicolour (tuxedo pattern) cat with intense sapphire blue eyes was found on a country road in Windellama, New South Wales, Australia by A M Schnieder. Since the Ojos Azules has not been imported into Australia, this represents a spontaneous mutation among the feral population. Whether it is the same as the American Ojos Azules mutation is not known.
Below: silver tabby female feral cat, black-and-white male, red tabby (with white bib) male, red tabby longhair male, odd-eyed tortie, black-and-white male.
Wain Harding, Secretary Egyptian Mau Breeders & Fanciers Association (May 1971 Cats Magazine): “I was very interested in your recent article ‘The Cat That Couldn’t Be Bred’ by Mrs. Sayer. The Egyptian Mau breeders have recently had four blue-eyed kittens from different parents. These kittens are beautiful well spotted, silver Mau with deep blue eyes. All of these kittens share a common grand or great-grand parent (Kattiwyckes Gulliver x Kattiwyckes Trinket owned by Mrs. Ann Cahill). The first of these blue-eyed beauties appeared in one of my own litters (Kattiwyckes Umm Usada bred to her son Bast’s Zosser). I kept waiting for this kittens eyes to change as it is often four months before a Mau kitten’s eyes change from blue to the desired gold. I exhibited Chui Chali at the Fall Baltimore CFA show when he was six months old. I still insisted that his eyes would change to the admiring public. The kitten’s eyes never changed and he grew up and sired a litter now nine days old. We are anxiously waiting for the kittens to mature.
The second kitten appeared in a litter of Ann Cahill’s breeding. This beautiful young female, Kattiwyckes Dynamite, will soon be bred to Bast’s Chui in the hopes of producing an all blue-eyed litter. The third blue-eyed baby appeared in a litter of kittens bred by Barbara Abruzzo of New Jersey. The fourth kitten appeared in a litter bred by Joan Dewberry of Ta-Mera Cattery in California. We are very proud of our blue-eyed stock and have great expectations for these kittens. The most plausible explanation for the blue-eyed mutation has to do with the amount of inbreeding that has been necessary to keep the Egyptian Mau the pure, natural breed it was upon importation. Bast Chui Chali is now at stud to Egyptian Mau queens of pure ancestry as well as to approved Foreign Short- hair breeders interested in breeding a foreign type cat with blue eyes. We are helping a breeder develop blue-eyed blacks at the present time. The Egyptian Mau Breeders and Fanciers Association are happy to announce blue eyed Mau kittens and gene-carriers in the near future.”
In Cats Magazine. October 1972, Mrs. Dagmar Thies, from W. Germany wrote: “From an American cat-breeder, I recently heard that it would be possible to breed blue-eyed Black Foreign Shorthairs. We are breeding these wonderfully charming cats, Calling them “Ebonies.” After three generations of back-crossing self-coloureds to best typed Siamese we started mating Black to Black, and now we have the third generation of Ebonies. Two of our queens, one second and one third generation self-bred, are showing very light blue-green eye colour, without any yellowish glimmer. Would it be possible to reach blue-eyed kittens by back-crossing them to Siamese once more? Do you think that blue eyed Blacks would be worth breeding?”
Don Shaw (Cats Magazine’s resident geneticist) answered: “Your letter is most informative. The enclosed photographs were excellent. You have truly obtained, the Foreign Shorthair type in your Ebonies. In answer to your question; there is no genetic evidence to my knowledge which would suggest that the genic factors for blue-eyedness in self-colored cats can be obtained by crosses with Siamese. All the data thus far examined by us, indicate that the basic causative genic factor for blue eyedness in Siamese is intrinsic to the “cs” allele which is responsible for the temperature sensitivity color-point affect. If this is indeed the case, then this genic factor must be eliminated if you hope to obtain true breeding self-colored cats. From the information at hand, it is very likely that the two queens you described may have picked up eye color factors from the self-colored cats in their background which may or may not be being amplified by the “cs” allele, which could still be heterozygous in these particular, cats. No, I do not think that crosses with Siamese would either help you toward blue-eyed black nor would it help you toward your goal of true breeding Ebonies. Are blue-eyed Blacks worth breeding? Yes, from a genetic point of view they would provide us with entirely new concepts concerning eye color. Over the years I have heard reports of truly blue-eyed Blacks, but in my ten years of searching I have not come up with one valid case. Somehow, people say they have had them or have seen them, but when I go to examine the cat it is not there, or the eyes are green. They may be bluish-green, but nonetheless they are green, not truly blue as we get in Siamese and Blue-Eyed Whites. From a sales view point, I can assure you there would be a market in this country and I strongly suspect the market would be world-wide.”
In Cats Magazine July 1973, Elaine Brenna wrote: I read with interest your remarks in the March [issue] about the rarity of blue-eyed black cats. I am not a breeder of cats although I have two, and a picture of one is enclosed. As you can see Tosca's eyes are quite blue; there is no camera distortion here. He is also black, but not quite jet black. This picture was taken when he was about four months old; he's now six months old and has matured quite a bit. He has not been altered. His eye color is still the same and his coat has taken on a glossy, sleek appearance. He resembles a Burmese quite strongly except for his eye and coat color. Tosca is quite intelligent and makes a wide variety of sounds. He does have one white patch on his lower abdomen. Whether your article was referring to Blue-Eyed Persians that are black or just blue-eyed black cats in general I don't know. I thought I'd send this on to you for information purposes only. Tosca's origin is unknown.”
Don Shaw (Cats Magazine’s resident geneticist) answered: We greatly appreciated the information and picture of the "blue-eyed black." Unfortunately, we cannot reproduce the picture, but according to the photograph, Tosca does indeed have blue-eyes. In fact, they are a fantastic blue. If, as you say, there is no camera distortion, they might be called royal blue - it is virtually unbelievable! The closest thing to this eye color I have ever seen is a deep violet blue which can be traced back to a particular Siamese line. A similar deep blue is relatively common in very young kittens right after they open their eyes, but this generally changes by four months and certainly by six months. If they are still this fantastic blue at eight months, we at CATS would seriously consider publishing a color photograph if you could obtain a good sharp color transparency with neutral background. That blue-eye coloring would be overwhelming on a cat of any color. >Now to the genetics — you state that there is “one white patch on his lower abdomen: "This would indicate that he is carrying the pied, white-spotting S-allele. Am I seeing things, or is, it camera distortion, or is there really a tiny cluster of white hair at the inner edge of the left eye near the bridge of the nose? Also please examine his toe nails—are they all showing darkish marrow? If this blue-eye coloring remains, you may well have an excellent example of the S-allele's producing blue-eye with very little indication of its manifestation in producing white-spotting.”
CONSIDERATIONS FOR ALL BLUE-EYED BREEDS
There are various known and unknown genes involved in white spotting and in blue-eyes. In solid white and high white cats there is a link to deafness because melanin is involved in the development of the inner ear (the part that detects sound vibrations). BAER testing (acoustically evoked brain stem responses) is used by many European breeders to ensure that they don't deliberately breed deaf cats. The prevalence of deafness and partial hearing in an experimental colony of white cats was 67% (deaf - 0.55 coefficient of heritability) and 29% (partial hearing - 0.75 coefficient of heritability) which suggests a pleiotropic major gene (a gene that has 2 seemingly unrelated traits) and the likelihood of polygenes.
Geigy CA, Heid S, Steffen F, Danielson K, Jaggy A, Gaillard C (2007). "Does a pleiotropic gene explain deafness and blue irises in white cats?". Veterinary Journal. 173 (3): 548–553. PMID 16956778.
This was commented upon by Strain GM, in "Deafness in blue-eyed white cats: the uphill road to solving polygenic disorders."Vet J. 2007 May;173(3):471-2. Epub 2007 Feb 21.
In some species, such as ferrets and "splashed white" horses, certain white spotting patterns are linked to mild forms of Waardenburg syndrome (syndrome means "collection of traits consistently occurring together"). Though suspected by some owners, this has not been confirmed in white-marked cats. Mutations in several different genes can cause similar visual appearance. The traits occurring together include a specific distribution of white, premature greying, brilliant blue eye-colour in one or both eyes, or eyes where the iris has two different colours (sectoral heterochromia), congenital deafness, broader nose and shortened tail. Apart from deafness, which is not unusual in blue-eyed white cats, the traits are cosmetic and have less impact than (for example) skeletal problems in Scottish Folds or spinal problems in Manx, which are both managed by careful selective breeding.