CAF Federation

The Genetics of Caramel and Apricot Inheritance

This article is reproduced by kind permission of club member Mrs Di Clarke, who has been granted permission by original author
Hetty Berntrop to use the article wherever she feels it may be appropriate. It certainly has its place here!!

Caramel is a subject of controversies in some Cat Fancies.  So I better start with a little bit of history. Although some people think caramel and apricot are new colours, caramel has first been identified as a different colour in a litter in England as far back as 1974.
One of the kittens was what we now call a caramel! The first apricot was registered in 1975.
The mother of the caramel  was the result of a cross of a Shaded Silver dam and a Siamese Red point sire carrying dilution (blue). The poor typed Shaded Silver dam came from a mismating between a Chinchilla Persian and a Siamese.
The caramel cat was Scintasilva Sue (sometimes referred to as Scinta Silver Sue).
Pat Turner rescued her mother from the owner. As soon as the litter was born, it was reported that the kittens were silver. Pat Turner looked at them and decided she would like to breed silvers and use these cats.
Nobody knew much about silver in these days. The mother of Scintasilva Sue was mated to a lilac point Siamese and produced another all silver litter.

A daughter of Scintasilva Sue, Scintilla Serene Sunset (a tortie shaded silver Oriental) was mated to Southview Trappist, a chocolate longhaired Oriental cat. He has been used for the foundation of the Angora's (Oriental Longhaired Cats) in the UK. He carried cinnamon and recessive white (or blue-eyed albino: ca). From this litter came a kitten with a strange colour, a colour that nobody had ever seen.
This mating had been done deliberately to get information about the difference between unsound coats in Oriental chocolates and silverwhite roots in smoke cats.

At first it was thought that this colour was the result of the working of the cinnamon gene and the blue-eyed albino gene.  However, some time later after a lot of research of the pedigrees of the cats involved and after a lot of test matings had been carried out, it became clear that this was not the case.
It was realised that this was a totally new colour. Now several interested breeders started matings to find out more about this new colour and to decide whether this was indeed a new gene, whether this gene was recessive or dominant, how it worked and if it was worth going on with it.

After a while it was decided not to continue breeding with cats which carried the recessive white ca gene, as this gene was undesirable in the Oriental and Siamese cats.
Scintilla Serene Sunset, who did not carry the recessive white gene, had been mated to two other males and had given litters, which included caramel kittens.
A lot of the silver Orientals and Siamese bred by Pat Turner kept on producing caramels.
A very good caramel was Scintilla Koffee Ole who went to a breeder of Oriental chocolates who used him quite a lot. This breeder also owned Gd Ch. Folklore Moonwolf, a black spotted oriental, who was also a descendant of Scintasilva Sue and he must have sired several caramels (as people reported), although there were hardly any judges at that time who could recognise this new colour. Moonwolf is at the back of many "Megrim" cats and has been extensively used because he sired such beautiful kittens.

Scintilla Serene Sunset was mated to a SP Siamese who carried blue and she produced a seal silver tabby point male, Scintilla Rosario, he was the father of Scintilla Koffee Kreme, a Siamese caramel point.

Scintilla Koffee Kreme mated Scintilla Sugar Icing, blue silver spotted Oriental, and she produced Scintilla Muted Mink, a caramel silver spotted Oriental. The type of the caramel Siamese and Orientals was improving all the time. A son of Muted Mink and Amoureuse Melody (Oriental black carrying blue), was Scintilla Pastel Royale, a caramel silver tabby point Siamese.

Pat kept using sons and improving type all the time. Scintilla Caramellian, chocolate tabby point Siamese, was the next in line. Pat Turner did most of this work on her own, as not many people were interested.
Unfortunately disaster struck her cattery and she lost a lot of her cats. Only a few of her own caramels survived such as Scintilla Caramelodot, a caramel spotted tabby.

The colour with which this new colour is most often confused with is lilac. If you compare a good lilac silver tabby and a caramel silver tabby there is a very distinct difference between the two, especially in daylight.
Caramel is more brownish with a blue haze or metallic tint.

In Pat Turner's home in Eastborne in the South of England geneticist Roy Robinson, Peter Dyte, Maureen Silson and Julia May all came to help and try to determine the genetic heritage of her breeding investigations.

From an unsuspected side caramel slipped in as well. When the first tabby point Siamese appeared they were seal tabby or chocolate tabby point. However, most tabby points come from a mating of Macji Dom Dija, a seal point Siamese x a Silver tabby moggie. Their daughter Tiggi, mated to Chancasta, a seal point Siamese gave birth in 1960 to Miss Tee Kat, the well-known seal tabby point that can be found at the back of almost all Siamese tabby point pedigrees. The silver tabby moggie was found in the same street where a Chinchilla breeder lived. And in the old days cats were not couped up in cat runs and catteries all the time but often could roam free in the village?
So it really was no wonder that caramel popped up in Siamese tabby point lines too, although it took some time for most breeders to realise they had a different colour.

Spotlight Petit Burlinks, a seal tabby point grandson of Miss Tee Kat had a progeny of over 60 cats, mostly seal tabby, chocolate tabby and seal and chocolate points. So the gene that produced caramel went unobserved for a long time. And there were numerous other tabby points, which were mostly being bred with excellent seal points to get a better type and the tabby points became ever more popular.

Apricot, the colour that appears when a cream cat carries the dilute modifying gene, appeared as early as 1975 when Pat Turner mated Scintilla Serene Sunset to Taurus Kay Cavalier, a lilac point Siamese. Three kittens were registered by the GCCF as Apricot. Through the years more and more apricot kittens were born and they are now also recognised and registered as a different colour by the GCCF of the UK.

Somehow caramel must have gotten into Burmese lines as well. At the time Pat Turner had Scintilla Dresden Rosa, a caramel point Siamese born 16-08-1974 (out of Scintilla Serene Sunset by Southview Trappist, the chocolate Oriental longhair), a Burmese breeder came to visit Pat with a strange coloured Burmese. She came to ask Pat's advice about the colour, which appeared to be the same colour as Dresden Rosa. Both Pat Turner and Peter Dyte were convinced that this Burmese girl was caramel too. But she was registered by the breeder as a lilac to avoid any dissension in the Burmese breed about a new colour!

When I saw one of the first blue Burmese arrive in Eindhoven (Holland) years later, they had a distinct brownish tint on their backs. I was told all blue Burmese looked like that. I was surprised because I was used to blue point Siamese where a brown hue was not at all permitted. Looking back at this Burmese I suppose that amongst these blue Burmese there must have been a lot of blue-based caramel Burmese as well.

In the meantime Pat Turner had found out that the gene for caramel was dominant to blue and lilac and
that it was epistatic to red, black (seal) and chocolate (brown). Later on it was discovered that it was also epistatic to cinnamon.

This meant that the colours red, black, chocolate and cinnamon could mask this dominant gene and that they could be carried unobserved for many generations as long as there were no genes for dilution (blue) involved. Pat also found out that the caramel colour carried by one cat was sufficient to produce caramel, provided there were also two genes for dilution (blue) present. This meant that it must be a dominant gene.

Either you get a caramel cat or the cat is blue, lilac or fawn. You cannot have a blue, lilac or fawn cat carrying the gene for caramel. In the meantime a new name had to be found for this gene. First it was called Modifying dilute, but in the end Dilute modifier (Dm) has been accepted as the better name.
It is situated on a separate locus and can be inherited separate from genes for other colours, including blue.

The colour genes are always present in pairs: BBDD for a black cat, bbDD for a chocolate cat, BbDd for a black cat carrying chocolate and dilution (blue). Chocolate and cinnamon take the same place (locus) on the chromosome. This means that a black, blue or red cat can have one gene for chocolate OR one gene for cinnamon. If this cat had two genes for chocolate he would BE a chocolate cat. If he had two genes for dilution (blue) he would BE a blue cat; if he had one gene for chocolate and one gene for cinnamon he would also be a chocolate cat, because chocolate is dominant over cinnamon. (Although you cannot see this in the way the genes are written; they are both recessive to black).
The combination of two genes for dilution (or blue) and two genes for chocolate in a black cat produces another colour: lilac (bbdd). The combination of two genes for dilution (or blue) and two genes for cinnamon in a black cat produces the fawn cat (blbldd).
In the red cat two genes for dilution produces the cream cat. Chocolate genes have no visible effect in a red cat. Although the late Persian breeder Piet Pros? in Holland thought that red Persians carrying chocolate had a much brighter colour red.

The dilute modifying gene is dominant, which means that one single gene can alter a cat into a caramel, provided also two genes for dilution are apparent. This happens to blue, lilac and fawn cats that carry this gene.
In cream cats, which also have two genes for dilution, the extra Dm gene alters the cream cat into the apricot cat.
Cats with two Dm genes are homozygous for caramel, but can only produce caramel kittens when mated to blue, lilac, fawn and caramel cats or cats which carry the dilution gene.  
A caramel cat with one Dm gene (a heterozygous caramel) gives this gene to 50% of its offspring. A caramel cat with two genes for Dm gives one Dm gene to its entire offspring.
But you cannot see this gene in red, black, brown, seal, chocolate or cinnamon cats, because the dilute modifier gene (Dm) is epistatic to these colours. This simply means that the colours black (brown), chocolate, cinnamon and red mask the Dm gene and thus can carry the gene that produces caramel or apricot without showing it.

So these red, black, chocolate and cinnamon cats could carry the Dm gene for caramel or apricot along for many generations without anybody knowing it; as long as there are no genes for dilution (blue or cream) involved in breeding these cats, it will not appear.
And it can come as a big surprise when a black cat carrying caramel, undetected for generations, and also carrying blue, is mated to a blue cat and a caramel suddenly appears out of the blue!!

The dilute modifying gene modifies the colour blue, lilac or fawn cat into a caramel cat. Blue-based:
Blue changes to have a brownish tinge, darker than lilac, mud coloured almost. In tabby cats the caramel pattern gets a distinct metallic overlay, which can also be seen when kittens have some ghost markings they definitely look metallic too. There is another difference between cats that are blue-based caramels and lilac-based caramels. The blue-based caramels have taupe footpads and the lilac-based cats are cool toned dark lilac with chocolate brown overtones and chocolate brown hairs between their footpads when young.
The cats that are fawn-based caramels have a much warmer colour, reddish brown almost cinnamonlike with a soft blue hue and are very beautiful. Fawn-based Caramel Point cats are paler in colour but can easily be distinguished from Fawn Point cats.
Polygenes for warm and cold and for cool and warm also play a big role.
This means you can get different looking cats in the same colour range, just as you can get three or more different toned chocolates or blues or lilacs depending on the influence of their polygenes.
In my opinion the blue- and lilac-based solid caramel cats (and caramel points too), are a bit dull compared to other colours and by far not as attractive as tabby (point) caramel cats are.

In red cats the dilute modifying Dm gene changes nothing, but it changes cream cats to apricot: a hot cream with a metallic sheen instead of the powdered effect of the cream.
Caramel tortie point cats can be easily identified, as apricot is much hotter in colour than cream, while the caramel has that distinct metallic overlay, even in kittens. They look very attractive.
In breeding caramel pointed and caramel tabby pointed cats myself, I could only see a slight colour difference between my blue-based and lilac-based kittens. Each time I got almost the same shade of caramel. But that was probably due to the fact that I had cats which carried many polygenes for light and warm. Other breeders could sometimes make more distinction between these two shades of caramel.
Lilac point can be clearly distinguished from caramel point cats at an early age, as lilac developes only slowly, while Caramel pointed kittens developed their colour as quickly as blue or seal pointed kittens. When you have a very quickly developing lilac point you most certainly have a caramel point instead! The hairs between their toes are always more chocolate than lilac, also an indication for caramel.

Caramel tabby pointed kittens can be tricky if they also carry many polygenes for warm and pale colouring and especially if the cinnamon gene is also involved. Then it often takes more time for the caramel colour to develop than 3 months, as the cinnamon gene seems to slow the colour development down.
And the colour in a caramel point changes all the time; sometimes it is more bluish, a week later it may have changed to more brownish overtones, which can be very confusing. When cinnamon is also involved in Siamese you might not know what colour kitten you have at first but have to wait a little longer, as the cinnamon gene slows down the development of the colour in pointed cats.
A good help is the pad- and nose leather colour. Fawn points have pink nose leather and paw pads, while fawn-based caramel points have a soft mauve colour. The hairs between their pads usually give a very good indication of the colour of the cat.

As I already have said, I like caramel best in tabby patterns. The metallic sheen of caramel contrasts beautifully with the warm ground colour in tabbies. I can still remember a gorgeous oriental caramel classic tabby I had to judge in the U.K.

I have seen the caramel colour for the first time many years ago when visiting the English breeder and geneticist Patricia Turner in the South of England. They were called Oriental "Pastels" at the time. Pat Turner told me that the American geneticist Don Shaw called the caramel colour  "Barrington Brown".
He saw this colour for the first time in the USA in Chinchilla Persians. Pat Turner used American chinchilla cats imported to England in her breeding program to develop silver in her oriental cats. When this new colour showed up unexpectedly, Pat Turner and her friends carried out a lot of matings to understand the nature of this colour.
In some Tai-Bagheera cats in Germany, also bred with the help of American chinchilla cats (Jemari chinchilla's if I am not mistaken) to get silver Orientals, caramel popped up as well.

There was a meeting about caramel, organised by one of the independent Cat clubs in Rotterdam (Holland) in the late 80's (I am not sure of the year as I could not take all my papers with me to Australia). Pat Turner and many independent judges and also several breeders attended this meeting.
Pat Turner told us more about caramel and we discussed caramel intensively. There also the name "taupe" was used for the first time for the blue-based caramels. One of the Dutch breeders invented this name for her caramels as she thought it fitted the colour of her cats better than caramel. She had rather dark toned caramel cats (originating from the German Tai-Bagheera lines) and the meeting agreed that it was a good idea to call the blue-based cats with the Dm gene taupe and to keep the name caramel for the paler lilac-based cats with the Dm gene. 
We saw caramel, lilac, blue and taupe cats in solid and tabby Orientals and Siamese on this meeting. There had not been bred any no fawn-based caramels in Holland at that time.
Pat Turner promised to give the information about taupe and caramel to Roy Robinson, as she did. In Roy Robinson 3rd edition of Genetics for Cat Breeders he referred to taupe, but unfortunately for the lilac-based caramels instead of for the blue-based, as was the intention. 

When I bought the 4th ed. of Genetics for Cat Breeders and Veterinarians this year, with very interesting new facts about many things, I discovered that taupe was still used for lilac-based caramel, which was never the intention. Fortunately the GCCF of the UK never used the name "taupe", but kept using caramel for the cats with dilute modifying gene.

A caramel cat (aaBBddDm) x a homozygous chocolate cat (aabbDD) gives 100% chocolate kittens with a gene for dilution (blue); 50% of the kittens are caramel carriers (aabbDdDm).
When a caramel point Siamese (aaBBcscsddDm) is mated to a chocolate point Siamese who carries dilution (blue) (aaBBcscsDd), you can get the following kittens:

| aBcsd |
| aBcsDm |
| aBcsd |
| aBcsd |

| aBcsdDm |
| aBcsD |
| aBcsd |
| aBcsD |

aaBBcscsddDm = caramel point Siamese
aaBBcscsDdDm  = seal point Siamese with gene for caramel and dilution (blue)
aaBBcscsdd = homozygous blue point Siamese
aaBBcscsDd = seal point Siamese with gene for dilution (blue)

Although dominant, the Dm gene can only express itself when 2 genes for dilution (blue) are present.
If the caramel cat (aaBBddDm) is mated to a blue cat carrying chocolate (aaBbDD) we can make the following diagram:
Caramel female : aBdDm  and  aBd
Blue male carrying chocolate: aBd  and  abd

| aBBdd | aBbddDm |
| aBbdd | aBBddDm |

aaBBdd =  homozygous blue
aaBbdd =  blue carrying chocolate
aaBBddDm = caramel
aaBbddDm = caramel carrying chocolate

If the caramel also carries chocolate she is: aaBbddDm.
Mated to a homozygous chocolate, aabbDD, the result will be:

| aBbDd | aBbDdDm |
| abbDd | abbDdDm |

aabbDd = chocolate with gene for dilution (blue)
aaBbDd = black with genes for chocolate & dilution (blue)
aabbDdDm = chocolate with genes for dilution (blue) & caramel
aaBbDdDm = black with genes for chocolate, dilution (blue) & caramel

Similar checkerboard diagrams can be made for crosses between apricot cats.

The English GCCF Standard of Points (the first registering body to recognise caramel and apricot) describes the colours as follows in the GCCF standard of Points:

Oriental Apricot
Coat colour: A hot cream with a soft metallic sheen, which becomes more noticeable with maturity. Tabby markings "may be evident, especially in kittens and should not be penalised in an otherwise good cat.
Hair: coloured to the roots. No white hairs.
Nose leather, Eye Rims and Paw Pads: Pink
Withhold certificates of First Prize in Kittens:
1. Coat white at roots
2. General Oriental withholding faults.

Oriental Apricot Tabbies
Markings: hot cream with a soft metallic sheen.
Groundcolour: pale apricot.
Nose leather, Eye rims and Paw pads: Pink

Oriental Caramels
Cool toned brownish blue, coloured to the roots. No white hairs.
Nose leather, Eye Rims and Paw Pads: brownish blue
N.B.: In my view there should be made a difference between blue-based, lilac-based and fawn-based caramels as they look distinctly different in solid Orientals (see above)

Oriental Caramel Tabbies
Markings brownish blue markings  with a metallic sheen on a cool-tined beige agouti ground.
Ground colour: beige or cool-toned beige.
Nose leather: brownish blue or pink rimmed with brownish blue
Eye rims and Paw pads: brownish blue
N.B.: The caramel tabbies I have seen on shows or at breeders all had a warm groud colour and I think the standard should be altered in this respect.

It took a long time before the caramels where recognised by the GCCF of the United Kingdom. They were registered as lilac point, lilac tabby point and lilac oriental (and sometimes blue or chocolate point) although they looked quite different. At some point you had to look hard to find a real beautiful old fashioned lilac coloured Siamese! Some lilac tabby points looked like they were carrying cinnamon, and that is what we thought in the beginning.  We have some dedicated breeders to thank for the recognition of the caramels. Now we can try to breed the real lilac coloured cats again.

In February 1987 I imported a lilac point Siamese male to Holland. I knew the breeder and the kitten's parents. I got pictures and he looked absolutely beautiful. So I went to pick him up at the age of 3 months, together with a pure chocolate point Siamese girl from different parentage.
All went well, but it surprised me that at 6 months his colouring was as well developed as for an adult lilac point.

He went to shows and became an International Champion but was refused further titles because his colour changed so much. You must already know what happened: he was not a lilac but a caramel point!
My "lilac" point mated the CP girl and the result was 1 SP and 3 CP kittens! I could not believe my eyes!  Later I repeated the mating with the same result, again 1 SP and 3 CP kittens. They were all looked at by experienced judges and were really SP and CP. But then the father could not be a lilac!
My cats all lived in the house and the stud cats had their apartments outside in the cattery, so the matings were all controlled.

I sent 7-generation pedigrees from both cats and a lot of hair I clipped from my studs tail base, (I could not show him any longer because of his strange colour) to Roy Robinson and Patricia Turner with the question what colour cat I had.
The answer from Roy Robinson was:" He is most probably a blue based caramel", otherwise he could not have sired these kittens. He was then re-registered in Holland as a caramel point Siamese.
I had a lovely caramel tabby point from him; with my caramel tabby point I bred many caramel point and caramel tabby point Siamese kittens.

My caramel tabby point came from a pale coloured line from Springfield's Bambuli (a real milk chocolate line from the famous chocolate point Siamese Ch. Kimoki Dagmar) and she carried many polygenes for light and warm and her kittens always looked pale caramel.
In her litters I could always see the difference between caramel and lilac point kittens at about 6 weeks of age. Tabby point kittens proved to be more difficult, but when I looked at the hairs between their paw pads that colour almost always gave me the right indication of their true colour.

I am indebted to Pat Turner in teaching me so much about caramel and apricot. She was always prepared to answer all my questions without delay and give information about all the knowledge she had gathered and discovered and she still is a great friend. I think  that we in the Cat Fancy are really fortunate to have her amongst us!  Aside from the information she gathered about the inheritage of silver, she has also been busy in the development of the Foreign Whites.
The latest breed she developed is the Seychellois, which she did while in the C.A., associated with Fife.
Pat Turner did not invent caramel and apricot, as some people thought in the beginning, but she gave a name to the colours that originated in cats carrying this gene. She discovered the workings of this dilute modifying gene through a lot of matings and much hard work. Due to her health she stopped breeding and judging, but she is still very interested in the development of the colours she named. A big thank you Pat!

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