Non-equines and other tangents



Elvis Brown posted this very interesting Greyhound to the blog’s Facebook page, and was kind enough to give me permission to share him here. Mr. Brown says he owned the dog from age three, and that he had always had the white spots, though at that time he was black. He is thirteen in this picture, and the greying is due to age. Mr. Brown also mentioned that someone from the Greyhound society had previously seen an Irish-bred Greyhound with a similar pattern.

These are somewhat reminiscent of Tetrarch Spots – sometimes called chubari spotting – in horses. Those take their name from the famous Thoroughbred, The Tetrarch, who was well-known for the unusual white spots on his coat. His daughter Mumtaz Mahal and (to a lesser extent) granddaughter Mumtaz Begum were similarly marked.



This kind of spotting in horses is associated with progressive greying, but progressive greying in dogs is different. In dogs grey is strongly associated with black pigment (eumelanin). It is most often seen in longhaired breeds, like the Bearded Collie, or in breeds that do not shed, like the Poodle and the Bedlington Terrier. Greyed dogs tend to be lightest where their hair is the longest, like on the topknot of some of the terriers, and darkest where the hair is short, like on the ears. This suggest that the hair loses pigment as it grows longer, rather than with each shed like a horse. Another interesting aspect of greying in dogs is that while it lightens black pigment, if the gene for black masking is present, it does not alter the black there. That is why Kerry Blue Terriers are born black and turn blue-grey while their face remains dark. The black mask, which would not otherwise be visible on a black dog, is revealed by the greying.



Whatever caused the white ticking on this Greyhound, it does not sound like it is related to greying. In size and placement, the white spots actually look a bit more like Birdcatcher Spots, which are more common on red horses than black ones. Quite a few horse owners report those as increasing in number with age, so they could be considered progressive, though the various kinds of white ticking in horses is another under-researched topic. I have some images to post for that, as well as an update on a related horse from a previous post, for tomorrow.

I want to thank everyone for their patience. I have been called away to deal with family issues more than usual this year, and I apologize that I have once again gotten off-track. Barring any further complications, the current topic of color oddities will return tomorrow.

In the meantime I wanted to share a link to a really interesting dog. Recently a puppy was discovered with the same pattern as a siamese cat. This coloring is a form of partial albinism and is caused by a mutation of tyrosinase, one of the enzymes involved in producing pigment. The mutation makes the enzyme temperature sensitive, so that the warmer parts of the body are pale while the cooler areas (like the extremities) are dark. For that reason, pretty much any mammal with this type of mutation is going to look much like the cats. In addition to cats, the pattern is also found in rabbits, mice, guinea pigs and rats like the ones pictured above. (The photo was taken by Diane Özdamar, whose wonderful gallery of rat photos can be found here.)

Up until this one was found, there was no evidence that dogs had a similar mutation. The puppy was said to have been found on the streets of Russia, but he disappeared before he could be tested. A photo of him can be found here. I wish I knew more, but that is all the information I have. Perhaps there are more like him somewhere in Russia. It would be a shame to lose the color now that it has occurred.

In the previous post, I talked about the two things that cause horses to have small dark spots on a white background. The first was the leopard pattern (Leopard Complex + Pattern1) and the second was the homozygous tobiano pattern. In this post, I want to talk about how dark spots on a white background are different in dogs.

Dalmatian dogs look like leopard appaloosas. It’s the same white background and the same small, round spots of color.  But Dalmatians are genetically very different from leopards. In fact, they have a lot more in common with the homozygous tobianos. That’s because they are “pinto” dogs. They just happen to be missing (or at least mostly missing) their dark patches. In fact, if you can imagine someone starting out with a classic tobiano horse – dark head, large round areas of color on the body – you have a good idea of what the basic piebald pattern is in dogs. In fact, in some countries the name for tobiano and the name for this pattern in dogs is the same: plating. Plattenscheck, platenbont – plate pinto. It makes sense, since tobianos have large “plates” of color on a white background. These dogs do, too. Or at least they started out that way. Here is a popular sire of English Setters from a little over a century ago.

His pattern is very reminiscent of tobiano. But breeders did not care for the patches, so they began breeding away from them.

In dogs, this kind of pattern is often called “extreme piebald”. It is still a “pinto” dog, but it doesn’t have a lot of color left, even on the face. English Setter breeders were not alone in this preference. The Dalmatian breeders were selecting for the same thing. They did not want patches, or even dark ears. They wanted all-over round spots.

Those round spots, which are visible in all three of these English Setters, look a lot like cat tracks to someone familiar with tobiano. What makes them different is that they aren’t actually part of the plating pattern. They are a separate thing entirely. For English Setters and some of the other sporting breeds, that’s the “Belton” pattern. The more technical names for it – ticking and roaning – are unfortunately for us horse people, already taken by very different patterns. So for now we’ll just use Belton to avoid making this any more confusing.

Belton adds dark spots of color to the areas the piebald pattern leaves white. What dog breeders have done is manipulate the scale and spacing of those spots of color. All three dogs at the top of this post have what are believed to be variations on this kind of patterning. The English Setter to the left is of course the original Belton pattern. The Dalmatian in the middle is likewise has a Belton-type pattern, but he also has some kind of modifier that has made the spots larger, rounder and more distinct. (Some of the distinctive nature of his spots are, of course, because he is a sleek-coated dog compared to the setter.)  The Australian Cattle Dog at the end has a Belton-type pattern that was modified to the other end of the spectrum, with spots that have gotten smaller, less round and less distinct. In some breeds, this is what is called Roan. There is some debate about whether Roan and Ticking in dogs are truly separate, or just variations on the same gene. I am not aware of any papers yet published with molecular studies, but it does seem that roan dogs, when outcrossed to non-roan breeds, end up with offspring that look a lot like the Belton setters. Certainly whether these are separate, similar genes or the same gene with layers of modifiers, the end result is that dogs have independent factors that will “recolor” the area that a piebald gene left white.

It didn’t seem that horses had that, at least not until recently.

In 2009 a French sport horse, Vision Morinda was foaled.

Clicking on the image above will take you to the website for her breeder, and her page which has many high-quality photos of her at all ages.

At first glance, it is tempting to assume that Vision Morinda is a tobiano with very loud cat tracking. The problem is that she cannot be homozygous. Her dam is brown. (Note that the mare she is pictured with is a surrogate. Her dam, Scarlett Fontanel, is pictured here.) But perhaps even more intriguing is the fact that her spotting seems to have intensified as she matured. That’s something that is typical of the Belton patterns. As most people are aware thanks to the Disney movie, Dalmatian puppies are born white and develop their spots later. That’s true of the English Setters and the Australian Cattle Dogs. Here is my friend Mary’s (extremely cute) Cattle Dog mix, Volt, as a puppy. (Thank you, Mary, for letting me share your photos!)

As you can see, he looks like a white dog with black patches. He is an extreme piebald. That’s why he has white ears. Well, mostly white ears. He was already starting to show some spotting there. His back and sides, however, looked white. But here is Volt today, as a grown dog.

As you can see, he developed his ticking – the Belton-type pattern – over time.

In a less dramatic fashion, Vision Morinda seems to have spotting that intensified as she matured. (Her breeders even comment on her page about the surprise of getting an English Setter color on their horse.) The spotting on her is also different, visually, from a typical tobiano with cat tracks. The pattern is evenly distributed. The spacing does change somewhat (notably across her shoulder) but it still is pretty consistent across the white areas, rather than clustering into spots or patches. It looks like the ticking you would see on a dog, not a horse.

This raises the question of whether there is some factor in horses that can add ticking – a Belton pattern, so to speak. I have a few more horses to share, all with odd spotting patterns. None are quite like Vision, but all have unexplained dark spots inside white patterns or markings. They all come from my “weird stuff” files. That’s where I put things that don’t make sense, or just seem “off” in some fashion. Sometimes enough of them accumulate – like the odd late greys from a few months ago – that it seems like there might be some thread connecting them all. I am not sure these horses really have a common thread, because they do have some visual differences, but I’m going to start posting them just to see if more turn up. That’s what happened with those greys (I have more that I need to post in the future, by the way!) so maybe sharing them will bring others out of the woodwork!

(Images at the top of the post are courtesy of Wikipedia. Images of historical English Setters come from The Pointer and Setter in America, published in 1911, and Country Life, Volume 22, 1907.)

Images of leopard appaloosas with Dalmatian dogs are always eye-catching. Certainly they can look quite closely matched, like this Polish Malopolski and his buddy. Even so, the patterns in the two species are very different in terms of what is really happening to the pigment on the animal. That’s probably off in the weeds for most owners and breeders, but for artists the distinction is actually pretty important.

This touches on one of the reasons why artists who develop an interest in horse color often have such a different perspective. Usually the kind of information a breeder needs is predictive. That is, they need to know what might likely result from crossing this to that, or what they might need to cross if this particular end result is what was wanted. What artists need to know isn’t about prediction nearly so much as it is about possibilities. Not so much what might happen, but what could happen – even far-out-there, not-very-likely, could happen. That’s because artists often want to add something for interest or for composition. For those producing realistic art, that has to be done within the constraints of what is possible. It doesn’t necessarily have to be likely, but it does have to be possible. This unique perspective became apparent to me a number of years ago when I gave my first presentation on horse color. In the question and answer period afterwards, someone in the audience asked if a horse could be both dappled and fleabitten at the same time. It was clear that was not the sort of question my fellow presenter, Dr. Sponenberg, often heard. But it is precisely the kind of question that equine artists ask all the time. Scientists might not notice this kind of detail on an individual horse, but for someone who paints horses, this kind of information – does this happen with this? – has a lot of practical value.

So why do artists need to understand the process behind appaloosa patterns? Spotting is a useful tool, because it breaks up positive and negative space. It makes the horse more visually interesting. If you are particularly clever, it can be used to draw the eye in a way that works with the composition, or to hide flaws. But spotting doesn’t just happen anywhere. It follows rules, and those rules depend with what is happening with the pigment. Understanding the underlying mechanism makes it far less likely that you’ll add some interesting detail that isn’t realistic. When dealing with rare combinations of colors and patterns, it might be difficult to find a reference image to consult. Knowing the process can tell you if there is a reason to bother looking in the first place, because it tells you what is possible. (And when you wing it without a reference, the knowledge will make for more reliable guesses.)

“Trouble”, sculpted by Sarah Minkiewicz-Breunig and glazed by Lesli Kathman.
In the collection of Melissa Gaulding.

This is a ceramic collectible with the kind of spotting (often called cat tracking) seen in homozygous tobianos. It’s a really popular effect. In my normal job (the one I have when I am not trying to get a horse color book to press), I have produced quite a number of these. The problem comes when this gets confused with leopard spotting, and most especially what happens when leopard patterns are combined with the tobiano pattern. That brings us back to the image at the beginning of the post. These three images – the leopard, the Dalmatian, and my ceramic foal –  represent three very different scenarios in terms of the underlying process. I want to take each, one at a time, and explain how they are different despite looking so similar.

This is the typical nose-to-toes kind of leopard. Most people would think of this as a white horse with black spots that have been superimposed on top. That’s not really accurate. From a genetic standpoint, this kind of horse is a two-step process. First she has inherited a gene that progressively adds white hairs to the coat. Those hairs, over time, are going to produce the fairly distinctive pattern known as varnish roan. If the pony in my illustration just had that first gene, she would look like a black version of this pony.

That first gene, known as Leopard Complex, sets things up for leopard but it doesn’t make leopard patterns itself.

That happens when the horse inherits a separate patterning gene in addition to Leopard Complex. In this case, that patterning gene is called Pattern1. What Pattern1 does is take the white from Leopard Complex and amplifies and organizes it. So while our horse looks like she is white with spots, it is perhaps more helpful to think of her as a horse that was roan, but Pattern1 has now taken that mixture of white and dark hair  and reorganized it. Underneath the white hair, what that horse may look like is closer to this.

This is what the underlying skin looks like. She probably does have some truly white skin in the area where a blanket pattern would go. Pattern1 does amplify the white, after all. But under it all she isn’t really a white horse, at least not in the sense that most people would think of as true white. She is more like a roan horse that has been modified a bit. That’s why even nose-to-toes leopards have faces that are shaded much more like a grey than a cremello, because for the most part the face has dark skin, not pink skin. And that is why a pintaloosa looks like this:

The true white areas of the tobiano pattern cover over the appaloosa pattern. The spots from the leopard pattern don’t spread over onto the tobiano pattern because the process with Pattern1 isn’t “add dark spots to the white”, it is “organize the roan into spots.” So the spots don’t happen where the tobiano pattern already took all the roan away. Without the color there in the first place, Pattern1 has nothing to work with.

Of course, if we moved our tobiano pattern out a bit, encompassing more of the dark skin and butting it up close to the “blanket” skin, we could probably get something that looked a bit like the leopard spots migrated over some of the tobiano.

Even so, the spotting is still concentrated in such a way that shows it is an appaloosa pattern with a tobiano pattern layered over the top of it. The spots on the flanks might look like they are in the tobiano white, but really they are just in an area that was already white from the action of the Pattern1 gene. The action is still the same. The tobiano is there adding true, pink-skinned white on the horse, and underneath it Leopard Complex and Pattern1 are just doing their thing.

Even with the tobiano bumping up to the pink-skinned areas of the leopard pattern, it still looks different from the kind of spotting that comes from a horse having two copies of the tobiano gene.

This is a (presumably) homozygous tobiano with cat tracks. Whereas Leopard Complex is a roaning process that Pattern1 takes and organizes into the leopard pattern, this type of spotting is more like a not-entirely-successful attempt to add some more color to a horse that already has large patches of color. Unlike the existing spots, which are large and opaque, these new spots are small and vary in opacity. Some just come through in specks.

Cat tracking tends to cluster around the existing spots to some extent, almost as if these new spots want to occupy the same general area as the existing spots. This is quite different from the spotting on a leopard, which tends to be dispersed across the body.

The exception is the hooves. Tobianos with cat tracks often have a concentration of spots around the coronary band, often turning the hoof completely dark or nearly so.

There are spots on the legs, but typically they are not as numerous as the ones around the feet. The same is true for the face. This is the face that goes with these feet. He does have a few spots in his blaze, but they are not extensive.

So how is this different from the Dalmatian? Well he really is a white animal with colored spots added on top. In dogs, the gene for this is usually called Ticking, but since ticking means something different in horses, I am going to use the older English term for the pattern, which is belton. This post has run really long, so I’ll split that over into a second part. And why delve into the belton pattern in dogs? Because lately there has been a handful of horses that have turned up that just might have that kind of spotting. At the very least there are horses with dark spots inside their white markings that are not tobiano cat tracking and not leopard patterning. More on those will appear in the next post!

My oldest son has been struggling with freshman Biology, so the concept of evolution has been on my mind a lot lately. I have also been reminded that evolution applies, not just to organisms, but of points of view. Sometimes it can be easy to forget that you once held a different view – unless of course you are foolish enough to write it down for others to find later. I have been writing about horse color long enough now that I have had that happen!

The reminder of this came from my husband. Longtime readers of the blog have probably picked up on the fact that I have a strong interest in the issues surrounding genetic diversity. It is, as I have mentioned, one of the themes that runs through the upcoming books. My close friends could probably warn readers that it is a tempting soapbox for me, particularly when it comes to the topic of dogs. That was exactly what I was doing – standing on that soapbox – when my husband reminded me that I once held a very different point of view on the subject.

He has every reason to remember this, since there was a time when the topic came up often between us. My husband is a physicist working in the field of optics. When we first met, he was intrigued by the possibilities of using genetic algorithms to solve complex design problems. A genetic algorithm is a mathmatical tool that narrows down variables by “breeding” the possibilities until an optimal solution is found. He was experimenting with genetic algorthims, and I had recently bought my first Arabian mare after spending most of my teen years planning my future breeding program. I was more than happy to explain all the different inbreeding, linebreeding and outcrossing schemes breeders had developed over the years.

My husband used this diagram to illustrate a chapter on genetic algorithms in his book. My understanding of how he used the idea doesn’t go much deeper!

I was familiar with them because I had my heart set on breeding animals, most specifically Arabian horses and Rough Collies. Since I could not convince my parents of the pressing need to start populating our home with dogs, or to acquire land for horses, I used the time to learn all I could for the day when I could do those things. When the time came, I was determined to be the most informed breeder possible. I devoured issues of Arabian Horse World. It was the 1980s and  the market for Arabians was at its peak, so each issue brought countless images for a horse crazy girl trying to determine just what qualities she would emphasize in her hypothetical breeding program. While other girls were pouring over fashion magazines and beginning to notice boys, I was filling ring binders with notes on bloodlines and affixing sticky notes to the important pages. As you can see, many of them are still there today.

I was smitten by pictures of the stallion *El Shaklan. The more something looked like an imaginary elven horse, the better. I was not an especially practical kid.

Arabians appealed to my artistic sensibilities. My interest in Collies came about in a more personal way. My grandfather was a Collie man, and encouraged me to read Albert Peyson Terhune’s books. Mr. Terhune had lived in the same town, and traveled in many of the same dog circles, as my grandfather’s family had when he was a boy. Like so many, I fell in love with the breed as it was portrayed in Mr. Terhune’s books. When my parents offered to give me my very own dog for my twelfth birthday, getting a Collie seemed a natural choice.

My grandfather’s favorite Collie, Glengay Sandy Boy. The inscription on the back of his photo is the Terhune quote, “a thoroughbred in body and in soul.”

I knew the “proper” way to obtain a quality dog. My parents, however, had different ideas. I wanted the perfect bitch with which to start my grand breeding program. My parents wanted something within their price range and a reasonable driving distance. The result was the dog at the top of this post. She came from a local farmer who raised a few Collies on the side. She wasn’t the potential foundation female I would have liked to have gotten, but she was a dog of my very own. When you are twelve, that counts for a lot.

My grandfather, when he received her photos, was quite critical. She had a pronounced stop, which was not proper. The angles of her face were all wrong, and she carried her tail in something awful close to a curl. But the real deal-breaker was her prick ears. As soon as she left the puppy stage and those ears went up, he ceased to consider her a Collie. Purebred Collies had tipped ears. She was, he insisted, nothing more than a “Whiffle Hound.” She was no relative of his beloved childhood Collie, Sandy.

Looking back, perhaps that reaction planted the seeds of doubt about what was valued in the animal fancies. It was obvious to anyone familiar with the standard that Brandy was a Collie of inferior type. I would have readily admitted as much. She was, however, a wonderful companion for a young girl. I thought she hung the moon, and her over-large, erect ears seemed like such a little thing in comparison to all that was great about her. We competed in obedience for much of her youth and mine, and it was pretty clear which of us did the better job. (Her ears were less of a limiter than my tendency to confuse left and right.)

Pictures of my grandfather with his ‘thoroughbred’ Collie, Sandy, and me with my ‘wiffle hound’, Brandy. Interestingly enough, we are the same age in these photos and adopting quite similar poses as we encouraged our dogs to do tricks for the camera.

But when Brandy passed away at fifteen years of age, I was explaining close linebreeding (the word I used at the time) to my husband. I had absorbed those ideas from the cultures that surrounded Arabians and Collies, and if anything was going to marinate a young animal lover in the twin concepts of “blood purity” and the usefulness of inbreeding, it would be the world of those two breeds.

But something happened along the way. Soon after we married, my husband’s work took us away from our small farm in Alabama, and to city life here in Charlotte.  I had already begun to question the consuming nature of raising dogs and horses, having seen that more closely through the experience of a number of friends. My interest in breeding animals became more academic, and less about laying the foundation for future activities. My mindset shifted from future breeder to a person who would own a series of beloved family companions, and who just happened to be very interested in the topic of genetics and breeding.

It was the academic interest in animal breeding that exposed me to a new way of thinking about breeding programs. My interest in color motivated me to read journal articles. Many of the authors of those articles also wrote about breed conservation and genetic diversity. Over time, the ideas presented in those papers brought about an evolution in my thinking. Like classical evolution, the change was gradual – a shifted position here, and new insight there, until something quite different took the place of what had been before.

Thankfully for me, this process occurred in the peace and isolation of my own research. I could think about the issues involved in a fairly objective, unemotional manner because no one was clamoring for me to reach a specific conclusion. Now many of those issues have become a source of controversy and bitter battles within the animal fancies. This has played out most visibly in the British dog show world, which was rocked three years ago by the documentary Pedigree Dogs Exposed. That program brought genetic diversity out of the realm of academic papers and into the public square. It is pretty safe to say that unemotional is not a feature of the situation!

In this reaction, many have called the focus on genetic diversity a passing fashion. That is not surprising, since fashion has long driven animal fancies. That is a natural frame for viewing motivations behind breeding decisions. I do believe, however, that calling the subject a fashion does a disservice to kind of thought that goes into breeding decisions. The acceptable angles of a Collie head, and the proper carriage of their ears, is a fashion. The acceptable amount of white on the legs and face on a horse are a fashion. The benefits and hazards of an increasingly homozygous population is a much larger issue. The truth is that breeders are dealing with competing concerns, with uniformity and predictibility being the very essence of selective breeding, and with its opposite, heterogenity, being so closely tied to health. That is not fashion: that is about the limits of the system as it really is. It is essential to understand those trade-offs if breeders are to make good decisions.

As I have said, the Equine Tapestry books touch on the subject of genetic diversity. Because the way a registry defines “breed” and “purity” has a huge impact on colors – both what is ruled in and what is ruled out – it is quite relevant. It is also true that misconceptions about purity and breed integrity have big implications for animal coloring. Because color is so easy to see, and is often the result of far more straight-forward inheritance than things like conformation or breed type, it tends to be on the front line of selective pressures. But beyond those issues, it is my hope that the books will raise questions about how we integrate our growing understanding of genetics into real world breeding decisions.

I have a few more posts to make that expand on some of these ideas, since I want to bring some of these ideas back to issues that touch more directly on color. I also plan to add another page (much like the Splashed White Project page) to the blog with a collection of links for further reading on genetic diversity. Perhaps they can plant the seeds for a more rational dialogue on the topic.

Twenty years ago, when I talked to my husband about linebreeding, I never mentioned genetic diversity. It was not a concept that had come up in my reading at that point. Today when I pulled the copy of my husband’s book, thinking I might find some of the images that came from his work in genetic algorithms, I found the following passage:

We intuitively know that larger populations will bring greater diversity and better sample the solution space. If the ranking function is nearly flat, poor attributes will stay in the population longer. If the ranking function is steep, the population swiftly becomes inbred. A lot of mutation can slow the convergence, while no mutation will lead to premature stagnation through inbreeding.

That passage pretty much sums up the situation facing breeders. If the selection process (what he calls a ‘ranking function’) is minimal, we will get a mix of good and bad attributes and not a lot of control over which we get from any particular breeding. That is the part I knew. At the other end, if we use extremely strict selection, we run the risk of a dead end where the variables present are too limited to provide the answer to a problem. For many, my younger self included, that is the missing piece. I may not have understood then, but the math certainly gave my husband a more complete picture. It almost makes me wish I hadn’t spent so much of my high school algebra classes doodling horses in the margins of my notebook.

The last few posts about silver in horses, and merle in dogs, dealt with mutations that alter black pigment without changing red pigment. Those two pigments – red and black – are pretty straightforward in horses. In dogs, though, the term “red” can lead to confusion.

That is because red is used in some breeds, like Australian Shepherds and Dobermans, to refer to what is really an alternate form of black pigment. The same color is sometimes called chocolate (Labradors, Cocker Spaniels), liver (English Setters, Pointers), or brown (Newfoundlands). Although they can appear red-brown in color, the pigment involved is a form of black rather than red. That is why a brown-and-tan dog will have two different shades of red-brown on their body. The brown-and-tan Kelpie pictured above is a good example of this. The darker areas of his coat correspond with the areas you would expect to be black on a black-and-tan dog, while the brighter, copper areas are the places you would expect to be tan. That’s because he carries the mutation that changes the black pigment (called eumelanin) to brown. Because red pigment (called pheomelanin) is not changed by the brown (b) mutation, his copper markings stay the same color. Were this dog not carrying two recessive brown genes (bb), he would be the more familiar black-and-tan.

Red merle are the same kind of color as the Kelpie, only with the merle gene added. Here is a red merle Catahoula Leopard.

Like the Kelpie, genetically he is a black-and-tan dog with the recessive brown (bb) mutation. He also has the merle mutation, which has merled the brown areas (which have a black pigment, or eumelanin, base) but left the tan (which have red pigment, or pheomelanin) alone.

The brown mutation also alters the pigment in the nose, paw pads, lips and eyes, so that the dog takes on a fairly monochromatic brown appearance. Here is a darker brown German Wirehaired Pointer showing how the nose leather is changed. The dog behind him, although somewhat out of focus, shows that the lips are changed as well.

Brown dogs can vary a good bit when it comes to shade. Some are redder than others, while some are a lot closer to black. With darker brown dogs, like this Dalmation and this German Shorthair, it is perhaps easier to imagine that the brown color is really an alteration of black.

So far, all the dogs posted have been genetically black (or black-and-tan) with the brown mutation. Brown, unlike the silver dilution in horses, does change genetically red dogs, too. It doesn’t change their fur, which is red, but it does change their noses, paw pads, lips and eyes. The extreme piebald Ibizan Hound posted a few days ago is a red dog with the brown dilution.

See how his patches are more similar in color to the tan markings on the brown-and-tan dogs? That is red pigmented fur. His nose, lips and area around his eyes are pinkish because the brown (bb) changed what would normally be black to brown. Nova Scotia Duck Tollers are another breed that is genetically red with the brown mutation. They also have pinkish-brown leathers and paler eyes.

Contrast the nose and eyes with the typical Golden Retriever, and that is how brown changes a red-pigmented dog.

And finally, one more bit that tends to cause confusion with the term red in dogs. The Golden Retriever pictured above is a genetically red dog, but most people would not readily call that color red, either. Most genetically red dogs are actually yellow in appearance. That is why, when speaking of dogs, pheomelanin is sometimes called “red/yellow” pigment. In horses, that is not typically used. There diluted red often does look yellow, but that is not common enough that the term needs to be added. In dogs it can help to clarify what it meant by red – especially given the confusion with brown.

(The Kelpie, Catahoula and Golden Retriever pictures are all courtesy of Wikimedia Commons.)

The recent posts on eye defects in dogs reminds me that I meant to share a recent paper on eye defects in silver horses. The issue of eye defects first came to light within the Rocky Mountain Horse breed, where it was initially called Anterior Segment Dysgenesis (ASD). That name was recently discarded in favor of Multiple Congenital Ocular Anomalies (MCOA).

Since its discovery, questions remained about whether or not MCOA was directly linked to the silver dilution, or if it was a more recent mutation tied to one of the Rocky Mountain founders. The recent study, “Multiple congenital ocular anomalies in Icelandic horses“, tied the issue directly to the silver mutation.

In this study we have shown that the MCOA syndrome is segregating with the PMEL17 mutation in the Icelandic Horse population. This makes the hypothesis that the MCOA mutation has recently arisen unlikely.

The Icelandic population is significant because it has been isolated from other domestic horses since 982 AD. If silver Icelandics have the same problem as silver Rocky Mountain Horses, then it is far more likely that the silver mutation is involved.

One of the things that makes this interesting is that the silver dilution – which is linked to eye defects in horses – occurs in the same location as the merling gene in dogs. Both are  PMEL17 (SILV) mutations. Both also dilute black, but not red, pigment. Those are interesting parallels between two colors that have such a visually different appearance.

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