Colours of the Rainbow

Colour me Beautiful ... Colour me HAVANESE !
Coat Colour Inheritance in the Havanese

Havanese are uniquely different from other breeds in many ways. One of the most intriguing of these is coat colour. Few, if any other breeds, carry such a wide range of colour shades and patterns in their coats as the Havanese. Havanese are renowned as a breed which carries a large majority (some say all) of the colours and variants possible in dogdom. They come in black, blue, silver, chocolate, brindle, sable, red, gold, champagne, cream and white displayed as a single solid colour or in assorted combinations. Not only is this vast array possible, the effects of an assortment of modifying genes result in endless variations. The possibilities are mind boggling. There is probably no area of the breed which rivals colour for intrigue and confusion.

Some people refer to me as the "Colour Guru". Flattering as that is, I must dispel the notion. I am not a guru of anything at all, just a breed fancier with an insatiable appetite for information. I must admit though, that Havanese colours and the genetics behind them intrigue me above all.

If your chosen breed is a Maltese; it's a given that your puppy will be white. White sire + white dam = white puppies. That one is easy to understand. Now, let's move on to the Havanese. Two parents of different colours may produce 5 puppies in a litter, each a different colour from themselves and from their parents. As one breeder put it "Havanese litters can look a basket of Easter Eggs". No wonder confusion reigns. It's easy to understand why so many people believe that Havanese colours are erratic and completely unpredictable. Many people understand the basics of genetics, but once the talk turns to alleles, locus, series, expression, partial dominance, recessives, modifiers, polygenic traits, etc, our understanding gets a lot foggier. So, let's start with the basics.

Chromosomes ( Figure 1) are the structures that hold genes. Genes are the instructions. There are many different genes to blueprint every part of a living being (including colour).Individual genes each have a specific spot on the chromosome that is its home. Homebase for a gene is called a gene locus. Next we need to know about Alleles. Those are the different expressions of a gene Each gene may have a few variations (alleles) or many, arranged according to an order of dominance. For example: on the C locus - [C] is dominant over all the other alleles that follow; [c^ch] is second in line. It is recessive to [C] but still dominant over [c^e] and so on. Even though a locus may have several allele possibilities, each living being only gets two, one from Mom and one from Dad. There are at least 10 different genes (likely more) that control colour in the Havanese. Different genes control different parts of the coat colour. Some genes make the different colour pigments while other genes control the distribution of these pigments within the individual hairs and all over the entire body.

Dogs have two kinds of pigment possible in their coats; dark pigment called "Eumelanin" and light pigment called "Phaeomelanin". In the world of genetics, any dog that has a predominantly eumelanin (dark) pigmented coat is referred to as "black", whether the coat is black, brown, charcoal blue or grey. In the same way, any dog that has a predominantly phaeomelanin (light) pigmented coat is referred to as "yellow" even though the actual colour of the dog may be red, gold, tan, champagne, or cream. Some genes affect just the dark pigment, others just the light pigment and some affect both. We also need to know about phenotypes and genotypes. A phenotype is what the dog looks like in appearance, while genotype is the non visible genetic codes responsible for making what you see. In some cases, there may be more than one gene code that gives the same appearance. Think of it like math. The answer (phenotype) is 4. There can be many different formulas (genotypes) to get to that same answer; 2+2, 1+3 , 5-1 and so on.

Each gene has a name designation. The ones we will look at are A, B, C, D, E, G, M, S, V, T and controversial K.

In simple dominance; if there are two dominant genes, the dominant is expressed. If there is one dominant and one recessive, the dominant is again expressed and the recessive will be hidden. The recessive gene will only be expressed if there are two recessives. In cases of incomplete or partial dominance, the dominant only partially covers rather than totally obscuring the recessive. These are the genes that cause such a shade variety and range of blended expression. In naming alleles, capital letters are used for the ones considered dominant and lower case letters are used for ones considered recessive The letter itself tells you which locus it is found on; for example, anything called [E] or [e] belongs on the E locus.Those little letters that are superscripted in the allele names are not nearly as confusing as they first appear and make sense if you look closely. They are variations which can be either dominant or recessive.If the superscript is attached to a capital letter, then it's a dominant variation and if attached to a lower case letter, then it's a recessive variation. Here are some examples.а a^t (tЕBlack and tan)аааa^y (yЕyellow)аа c^ch (chЕchinchilla)ааааааааc^e (eЕ extreme)аа k^br (brЕ brindle)а sⁱ(iЕirish Spotting)аs^p parti)

NOTE : There remains considerable doubt and confusion about the exact genes and alleles responsible for certain aspects of coat colour inheritance. The main controversies are "dominant black" traditionally put on the A locus and "brindle" traditionally put on the E locus. Till recently, researchers considered the Agouti Series (A) as being [A], [a^y],[ a^w], [a^s], [a^t], [a] and the Extension Series (E) as being [E^m], [E], [e^br], [e] Newer current research ( since 2003) indicates that dominant Black does not belong in the A series at all and that Brindle does not belong in the E series but rather that these two are part of a totally separate series which has been called K for ( blacK). This new information is what I am following. Do keep in mind that other sites may have Brindle on E and dominant Black on A.
Let's take a closer look at the genes and see what each one does and how it works.
The A locus - AGOUTI - this is the gene which controls dark pigment patterning and determines where and how the dark pigment made by E will be deposited. Agouti and Saddle variations are tentatively shown here ; we dont know for sure the correct order of dominance, but it is theorized that they may both belong between [a^y ] and [a^t ]. Dominance appears to be [ A^y ] >[ a^w ] > [ a^st ] > [ a^t ] > [a]. Lower case [a] is considered by some to be a recessive black ( not scientifically proven yet) so it has not been included here. To some extent the A series alleles appear to be incompletely dominant over each other. For instance; [A^y ] appears to be incompletely dominant over [a^t ], which may help to explain why some sable dogs appear to have tan points, or a tan mask. An [A^y a^t ] dog may appear as a Sable with Tan points . The Sable coat in a Sable/Tan is usually darker, having more black overlay and tips than a pure [A^y A^y] dog. In all likelihood, its not that simple and there may be other genes in play. To further complicate things, the A gene appears to have 2 separate and distinct parameters controlling its expression. One seems to control the colour of the head, neck and back (dorsal areas) and the other seems to control the chest, belly , legs (ventral areas).

[A^y] Sable patterning is considered the most dominant allele on the A locus in current genetic research. Sable is generally a predominantly light coat with dark tipping or overlay. Light coat colour can be two or more shadings of colour. [a^w] Agouti or Wild patterning is a banding of colour throughout each hair(dark/light/dark/light) - like a rabbit or wolf [a^st] Saddle tan lays the dark coat out in a saddle pattern. This may be a pattern on its own or may be an extreme variation of Black and Tan recessive [a^t] produces the black and tan pattern. It can be hidden by other A patterns.

phenotype ^{Dog Displays
Sable}

^{Dog Displays
Agouti}

^{Dog Displays
Saddle}

^{Dog is Black and
Tan}

genotype [ A^yA^y ] or [ A^ya^w ]
[ A^ya^st ] or [ A^ya^t ] [ a^wa^w ] or [ a^wa^st ]
[ a^wa^t ] [ a^sta^st ] or [ a^sta^t ] [ a^ta^t ]

The B locus - BROWN - determines what colour the dark pigment will be. Black or Brown. This affects the coat colour as well as eye colour and leather. The term "leather" includes nose, eye rims, lips and pads. Order of dominance
is BB > Bb > bb. True brown ( chocolate) cannot have any black pigment anywhere including eyes and nose. The same as black, chocolate can appear as the dark coat in any mixed dark/light coat. so there may be Chocolate Brindles, Chocolate Sables and Chocolate & Tans. If there is a chance meeting of two recessives [bb] and [ee], clear dogs with liver pigment may be the result.

B makes black pigment. b makes brown instead of black

phenotype Black

Brown ( Chocolate)

genotype BB or Bb bb

The C gene in most species is called ALBINO. Others refer to it at the CHINCHILLA gene or the CONCENTRATION gene. The C gene acts like a filter controlling the development and expression of the light coat pigment. The C gene is considered to mainly affect light pigment, though it may influence dark coat colour to some extent and is perhaps partially responsible for soft black, dusty darks and the like. The dominant CC does not filter at all, and allows a deep full intensity coat to develop. Filtered red develops less colour pigment and so shows as gold, apricot, champagne, cream etc. This may be modified further by a gene like plus/minus to make innumerable variations. Order of dominance is [C] > [c^ch] > [c^e] > [c]
Darker and lighter shades of each may blend into each other. For instance [Cc^e] and [c^chc^ch] can be indistinguishable. Note - Albino in mammals is the most extreme expression of this gene , fully recessive [cc] that prevents any colour from being made at all in coat, eyes and nose, and results in an albino animal with pink nose and red eyes (like an albino rat). Albino does not seem to appear to occur in dogs normally (though is possible) and is only mentioned here for interest sake. The white found in Havanese is actually extremely filtered [c^ec^e ] off-white. He does have some very watered colour, also has eye and nose pigment.

[CC] allows full deep intense colour to develop filters slightly resulting in a golden colour about half the red colour development is filtered the majority of the colour is filtered most of the red pigment production is filtered leaving the dog appearing white

phenotype
Red

Gold

Champagne

Cream

White

^genotype [CC] [C c^ch] [C c^e] or [c^chc^ch] [c^chc^e] [c^ec^e]

The D gene locus holds the DILUTION gene which dilutes all pigmentation made by B. DD will be full intensity with no dilution. This will allow a Black to be black and a Chocolate to be Chocolate. When dd acts on black, it results in a light silvery blue coat colour, with lighter eyes and smoky blue nose. Some refer to this as "Blue born blue" to distinguish it from the dark silver dogs that are called Blue . When dd acts on chocolate it results in a dilute chocolate ( Café-au-lait) colour with amber eyes and light rosy brown pigment. In some breeds , these dilutions are called Pearl, Isabella, Lilac and Mouse. There are few true dilute Havanese at this time, but occasionally they do occur. Order of dominance is [DD] > [Dd] > [dd]

Dominant [DD] gives intense colour to coat, nose and eyes with no dilution Recessive [dd] dilutes black affecting coat, leather and eyes. Recessive [dd] dilutes brown, affecting coat, leather and eyes

phenotype Dog is black

Dog appears silver Blue

Dog appears Café-au-Lait

genotype [DD] or [Dd] acting on black
[BB dd] or [Bb dd] acting on brown -
bb dd

The E locus (EXTENSION) controls production of dark pigment, deciding whether or not to make it and the extent to which it will be made. It can only deposit the colour it makes according to the directions of A and K. The order of dominance is [E^m ], [E], [e]. Em is dominant to E but "m"only affects a small area on the dog, and E expresses itself on the rest of the dog (according to A and K ) which is why you get masked Sables , masked brindles etc. Mask expression can range from covering the entire head to just raccoon eyes, or a dark muzzle. A face mask in some cases appears to also come in combination with dark paws and tail tip as well as a dark dorsal stripe ( this perhaps an interplay with other genes or modifiers).

Capital [E^m] is the most dominant, and deposits colour all over and includes a dark mask. Mask will not be visible on a solid dark dog but will show on other dark and mixed dark/light patterns. Capital [E] is also considered dominant, producing dark colour everywhere that A tells it to. There is no mask. [ee] is the most recessive, producing no dark pigment at all, even at birth.. Any amount of dark coat present, means it is a genetic sable not clear

phenotype Dark mask
Solid Dark dog - no mask

Clear dog

genotype(s) [E^mE^m] Or [ E^mE] or [E^m e] [EE] or [Ee] (carries clear) [ee] can cover any and all of the "A" series patterns

The G gene - GRAYING - does exactly what it says. It is often confused with the V gene but is not the same. While "V" filters colour development, "G" softens colour over time by greying the coat, much like premature graying hair in people. As graying progresses, coloured hair is replaced with silver, white or colourless hair. The more graying there is, the lighter the dog appears. Graying may start shortly after birth or much later. A dog with the graying gene will display a coat which has a gradual accumulation of white/colourless hair interspersed with the coloured hair. Incomplete dominance. Not all Havanese grey. Some do not.

dominant- Dog will progressively gray as it matures Dog will progressively gray
as it matures little or no graying

Phenotype

genotype [GG] [Gg] [gg]

The K locus (BLACK) is a somewhat newly formed series identified according to current emerging genetic research ( since 2003). Despite what most of us were taught, this locus is where dominant black appears to belong as well as brindle. There are at least 3 alleles in this series but there may be more. K controls dominant dark pigment. Full dominant black completely dominates all expression on the A locus. However it is still has to pay attention to the whims of the E locus. If no dark pigment is made [ee], then it has none to deposit. The order of dominance is [K] > [k^br] > [k].

Capital [K] is the most dominant, and deposits intense dark colour all over allowing no expression at all from the A locus. [k^br] deposits dark colour in a tiger striped pattern across the body . Brindle is generally considered dominant over all A patterns but it does appear to be incompletely dominant in regards to [a^t ] . See the colour pages for more information. recessive [k] is a normal expression of dark ( some call it recessive black expression). It does not dominate at all allowing the full expression of the A series pattens.

phenotype Dominant black
Brindle

Normal black expression

genotype(s) [KK]
[K k^br] (carries brindle)
[Kk] (carries normal expression) [k^br k^br]
[k^br k] (carries normal expression) [k k]

The M gene - MERLE - may or may not be a factor in Havanese. This has not been positively identified in Havanese to date but may be theoretically possible. Merle dilutes colour in patches creating a marbled, mottled appearance, also affecting nose pigment and eye colour. Doubled Merle genes in other breeds often produce health issues.

The S gene - SPOTTING - decides where the colour will be deposited and how much of the body will be covered with colour or be white. White is not a colour, it is an absence of colour. The spotting gene may make more sense if you think of white as a pattern, a mask that covers up the base colour of the dog. Dominance at this gene appears incomplete as possibly affected by modifying genes. More than one combination can look similar. Each one blends into the next. Note * - In many registries, any broken colour dogs are all called "Parti" regardless of how much or how little colour break there is.

All over solid colour with no white at all or only in minute amounts. Small amounts of white possible on chin, chest, feet and tail tip. Can be just a few hairs of white and may often be missed on light colour dogs. Irish Pied markings - coat is more than 50% coloured Close to 50% white and 50% colour. Some call this a heavy marked Parti. Classic Parti colour coat is more than 50% white with irregular splashes of colour Extreme Parti or Piebald - mostly white with minimal colour, perhaps just one or both ears

Phenotype

Genotype [SS] [S^si ] [sⁱ sⁱ ] or [S s^p] [sⁱ s^p ] or [S s ^e ] [s^p s^p ] or [sⁱ s^e ] [s^e s^e ]

The T gene - TICKING - decides whether or not the white areas of the coat ( determined by "S") will be flecked with
colour or not. A ticked coat may be referred to as Belton. Order of dominance is [TT] > [Tt] >[ tt]

[T] is dominant. It ticks all the white areas of the coat with flecks of colour and may be responsible for a heavily ticked coat while [Tt] still ticks but perhaps to a lesser degree. ^{[ t t ] is recessive. There is
no ticking.}

phenotype Dog is ticked
There is no ticking

genotype [TT] or [ Tt] [tt]

The V gene - SILVERING acts in a very similar manner as the C gene, but it acts as a filter limiting the development of dark coat colour( Black or Brown) Similarly it may also be affected by modifiers which produce a wide range of expression.
Where the Silvering Gene is affecting Black - Both Silvers and Silver Blues/Charcoals are born black with the colour generally clearing during the first year for a silver and 2-3 years for silver blue/charcoal. The Silvering gene only affects coat pigment. The nose pigment is black and the eyes dark. These are silvered expressions of dark colour ( Black) and can appear as the dark coat in any mixed dark/light coat so there may be Blue Brindle, Silver Brindle, Blue Sable, Silver Sable, Blue & Tan and Silver&Tan though these are certainly less common.

No filtering - allows black to develop fully One copy of the slivering gene appears to produces the soft charcoal colour that is often called Blue Two copies of the slivering gene produces the Silver dog

phenotype

genotype [B B] [v v] or [B b] [v v] [B B ] [v V] or [B b] [v V] [BB ] [V V ] or [B b] [VV]

Where the Silvering Gene is affecting Brown - Both Chocolate Silvers and Chocolate Blues(Mocha) are born chocolate with the colour clearing during the first year for a chocolate silver and 2-3 years for a chocolate blue. The Silvering gene only affects coat pigment. The nose pigment is Chocolate and the eyes medium/light brown. The Chocolate Blue and Chocolate Silver colours are silvered variations of dark (brown) and in theory could appear as the dark part of any of the mixed dark/light coat combinations. In reality, multiple recessives meeting other multiple recessives are very uncommon occurrences, though are genetically possible.

No filtering - allows chocolate to develop fully One copy of the slivering gene appears to produces the soft medium mocha brown colour. Two copies of the slivering gene produces the Chocolate Silver dog ( the colour of light coffee cream)

phenotype

genotype [b b] [v v] [b b] [v V] [b b] [V V ]

In an attempt to make it as understandable as possible, I have included a condensed Colour Chart as to what the different genes are and what they do. Each gene has a name designation. The ones we will look at are called A, B, C, D, E, G, M, S, T, and V. Chart does not have all the combinations possible in Havanese, but does show most of the basic ones.

The interplay of so many genes and possible variations is what makes Havanese colours so intriguing but also complicated; however, colour in Havanese is not as unpredictable as it first seems. Even only one litter can tell you a lot of information about the coat colour genetics of the Sire and Dam. Watching a litter as it grows to maturity can tell you even more. Let's now take a look at our Sample Litter

Occasionally we hear comments about "new" colours that are popping up in Havanese. The colours are not really new at all. They have always been there, but have just been hiding. They are very likely the result of recessive pairing. As there get to be more Havanese, the likelihood increases of a recessive allele meeting a matching recessive and getting the chance to be expressed. Remember: As Francis Greer said, "Genes don't go away, they hide."

Generally speaking, most Registries request that breeders register the puppies the colour they are at about 6-8 weeks of age. New changes in some registries ask that puppies be registered as the colour of the anticipated adult coat. This may cause much confusion as, in many cases, the colour may not develop as guessed. Also many colours lighten in the first few years but then darken again as the dog matures. Keep this in mind when researching pedigrees as the registered colours may provide little clue as to the actual genetic colours in the line. pedigrees are fun to have and intriguing to look at but keep in mind the limitations, as part of the information may be more a matter of interest than 100% genetically accurate.

This brief colour study is by no means complete nor does it show all possibilities; it is meant only as a basic overview of the genetics involved in Havanese colour. The information contained herein is colour inheritance as "I' come to understand it through my studies. As time goes by, changes may be made and definitions may become clearer as we learn more and more about the genetics behind coat colour. Anyone wanting to pursue more in depth study can do so as they wish.

It has been asked how its possible for the Havanese to have so many colours when the base breeds in its background and related breeds are white only, such as the Maltese, Bichon Frise etc. This is a very interesting question and one that bears looking at. Many of the breeds that are currently only accepted in white were not always historically so. See Maltese History for information on the history of coloured Maltese. The Lowchen and Bolonka which are related breeds do also come in many colours. In fact, a white coat in the Bolonka is undesirable. The Coton is permitted patches of colour though preferred white. Cotons with coloured patches usually lose that colour as they mature, very likely due to a strong chinchilla gene or a similar modifier. The Bolognese as well historically has colour in the background. We are told that the Bolognese is essentially white in color and must give an over-all white impression. This wording appears to indicate that other colours may be blended. One must also keep in mind that other breeds may have played a part in the development of the Havanese, such as one theory which links it to a small Poodle (see the Havanese History page). There is a rich history of colour behind the Havanese, some which we know and much which we do not.