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I think I need to work on final book edits every spring, because it seems to guarantee that a paper will come out within that time period. Animal Genetics has a short communication out with three new KIT mutations and one new PAX3 in horses. There are also two papers out on KIT patterns in dogs, which is new. Before this, the piebald patterns in dogs had been mapped to MITF, which in horses is the other site for splash white. The picture above is the German Shepherd that carried the de novo mutation. (I must give a special thank you to her owner for allowing me to include it here.) I had intended to put together a longer post on this pattern, known as panda, because it touches on the subject of blue eyes in KIT mutations. She obviously has blue eyes, and her owner confirmed for me that some of her descendants have had a blue eye or blue segments in their eyes. This follows the pattern that I have seen in the historical records of some suspected Dominant White horses (also presumed to be KIT mutations). Blue eyes do seem more common in the originators, and then appear to occur sporadically – often in a less pronounced degree – in the descendants. Interestingly enough, the MITF mutations are not associated with blue eyes. In fact, this family of Shepherds was the first instance I noted where blue eyes in dogs were linked with a form of white spotting so I was particularly happy to see the mutation formally identified.

What is interesting about the new equine discoveries is that they really do not fit neatly into existing naming categories. The KIT mutations have been assigned numbers in the “W” series, but at least one appears to be subtle white-booster rather than a true dominant white. I had been urged by a couple of researchers, as I got close to my publication date, to avoid the use of the term “sabino” and just use “white spotting”, and I see now why. I suspect this will become more complicated as time goes on and more mutations are identified. One thing does seem clear, and that is that several of these sites mutate often – apparently in ways both large and small!

Just as I did with the book last year, I’ll be updating the text to reflect this new information for the new full-color supplemental book. I had feared there might not be much new information, but this coupled with some of the things I have been researching in the last few months should make for a lot more content as well as more abundant (and colorful) images!

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This is probably the longest this blog has gone without a new post, and I want to thank all the readers who have stuck with me. I have not dropped off the face of the earth, but I have gotten sucked into alternate universe of preparing a book for press. I had hoped to continue posting here at least intermittently through that process, but juggling the demands of the book and the studio have consuming most of my time. If I am going to make my summer deadline for the new book, I am going to have to put the blog on hiatus for the next month or two.

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The upside is that the book that will come out this summer, Equine Tapestry: An Introduction to Colors and Patterns, has grown in scope. I originally intended it as the wished-for color version of the first volume, or more accurately, a color version of the first half which talks about the different colors and patterns. (The second half, which covers the individual breeds, had primarily black and white historical photos.) I also thought I could take the opportunity to expand some of the sections on the oddities, most of which were just mentioned in passing. I was particularly interested in expanding the entries on belton patterning, since I have come to believe there is more than one kind, as well as the various kinds of mismarks and somatic mutations. And of course, I could use more photos since color opened up the possibility to communicate so much more information. This opened the way for the “mission creep” that turned the original Equine Tapestry into a four-volume set! Well, now five volumes, since this new book is a supplement to the series.

It started with the belton patterning, and the fact that the outlines I used in Volume 1 were not really suited to showing the patterning. I needed a horse with a more clearly turned head to show the full face. I also thought that if I was going to do that, I might as well finish up the revised pony outline, too. The pony used in Volume 1 proved problematic because I drew him with long hair, making him a poor choice for any illustrations that communicated information about pattern outlines – which is most of them. So new outlines were drawn.

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The new outlines encouraged me to think about gaps in the information in the original book, and before I knew it I had plans for more charts, and a longer wish-list of photos to include – and much less time for managing the blog or even my personal correspondence.

My goal for the next few weeks is to draw some lines on the scope of this new book so I can accurately assess my publication deadlines. Once I have that done, I’ll better know when the blog is likely to go back online. I will still be posting intermittently on the blog’s Facebook page, since I can usually do that fairly quickly. (I am an avid user of my personal Facebook page, since that allows me to keep some contact with friends and family even in the most obsessive stages of book writing.) Rest assured, though, that it only seems like I am being silent! And once the book is put to bed, I will be chatting here again.

 

Charli

In the first volume of the Equine Tapestry series, I talked about two cases of unexpected dilute foals. The first was a Dutch Draft filly, Marinka van’t Heereind. The second was the Alt-Oldenburg filly Gaja. There is further information in the upcoming volume that covers the light breeds, where similar horses have been born to purebred Arabians. More recently, an entire family of Morgans has been documented that appears to have this same as-yet-unidentified dilution. Laura Behning has put numerous pictures up with photographic pedigrees on her Morgan Colors site. I highly recommend visiting her page!

Possible New Dilution in Morgans

There are also photos of the Arabian family with the similar dilution on the New Dilutions website.

Mireyenion Tos

The color has been called “light black” for lack of any better term, since it appears to dilute the black hair more strongly than the red hair. Because both the Morgans and the two Arabians are closely linebred, and because none of the parents are unusual in color, it is believed that this might be a recessive dilution. It should be noted that these horses have all tested negative for the known dilutions. In appearance, many have looked like the Laura Behning’s Morgan mare, Positively Charmed (“Charli”), who is pictured at the top of this post. Charli is a tested smoky black with the silver dilution. That particular combination produces a body color like milk chocolate, while the skin tends to have a purplish cast. Many of these horses also have paler eyes, but I have not yet seen a silver smoky that had eyes quite as pale as those seen in this Morgan family.

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In 1989, a horse I was riding lost his footing along a steep incline. The fall broke my wrist and shattered my humerus from elbow to shoulder. I was fortunate to have an employer that provided temporary disability, but confining a technical illustrator to a traction bed without the use of her dominant hand for several months is particularly cruel. A friend took pity on me, and suggested that I get a modem for my computer so that I could “chat” – one-handed, hunt-and-peck style – with people on the local computer “bulletin boards”. So by accident, I became an early adopter of online communication. Mostly I got to talk to young, nerdy guys about gaming (the old-fashioned kind, with polyhedral dice and character sheets) and computers. It would be three more years before the technology matured and spread enough that I could find people to talk about what I really loved, which was horses. Horses and their colors! Once that was possible, I filled countless hours chattering on that subject. (Some things do not change.)

One of my favorite topics at that time was pinto patterning, or more specifically, sabino patterning. Crop-out pintos were the focus of a lot of attention, and a number of articles and papers about “overo patterns” appeared around that time. I had also acquired complete, or nearly complete, sets of Walking Horse and Welsh Pony stud books, both of which included very detailed descriptions of markings. I began assembling data, using a numerical grading system based on the amount of white, trying to see if I could figure out why some horses ended up body-spotted. I became convinced that pinto patterns did not just pop up unexpectedly. I believed that the way we were looking at markings and patterns was mistaken, and that the horse world had drawn an artificial line between “pinto” and “not-pinto” – one that probably did not reflect the underlying genetics. Horses like the one below were universally recognized as pintos, but I was pretty sure you could take a lot of that white off and still have something that was genetically a pinto of some kind.

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I was so taken with this idea that I spent a lot of time online suggesting that maybe what we saw as markings were just part of a continuum that ended with a white horse. “Minimal” sabinos might look like an ordinary horse with ordinary markings. “Maximum” sabinos might be entirely white.

Having read a lot more papers (including some that others have kindly translated for me from their original Dutch or German), I know now that what I was suggesting was hardly new under the sun. Others had come to similar conclusions over the years. But it was rather unconventional thinking among horsemen at the time I began writing, and I got the message quite clearly from some that perhaps I was a little touched in the head. (I am sure that being overly enthusiastic about color, and sure of what I knew in a way that only seems to happen when you are twenty-something, did not help!)

At one point, in one of these online conversations, an older and more experienced horseman well-versed in Thoroughbreds finally asked, “Are you saying that all markings are really just patterns, minimally expressed?!” His implication was clear: this was crazy talk. I had not actually entertained that idea, but I had wanted people to consider that markings might occasionally work like a continuum. But his comment did make me think. Could that actually be true? Was it possible that all white markings were just minimal expressions of one of the patterns? I did not think that was the case even then, but it was an intriguing question that stuck with me over the years.

Nowadays, the concept of “minimum” and “maximum” patterns is pretty widely accepted. What I find interesting, however, is that taking that idea to its most extreme conclusion – that all white on the face and legs represent some minimum pattern – is also very widespread. What was once used to illustrate how ridiculous an idea might be, now has a certain following. In fact, if you spend enough time in forums and blog comments, you will soon find this “fact” being used as a verbal cudgel against someone “uninformed” about patterns. It seems we have come full circle in the world of online horse color discussions.

So is that the case? Are all white markings just patterns? Always?

The answer depends somewhat on how you want to define “pattern.” In genetics, piebald and “marked with white” are synonymous terms. Most horsemen, however, see a significant difference between a pattern (ie., something that may extend the white past acceptable or desirable levels) and markings (ie., something that will not extend past the extremities). Because that distinction can have serious implications in many breeds, perhaps putting it in those terms is most helpful. Does every horse with white on the legs or face have the potential to produce something body-spotted?

Or are there genes that, at their maximum expression, never extend past the lines that generally qualify  a horse in a pinto registry? Is is possible to produce this much white, say, and no more?

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What if it never goes even that far? What if you have a breed population where a horse like this represents the maximum end of the white patterning, even counting factors like base color?

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And it does look like there are marking genes that do not, in themselves, produce pinto patterns. The assumption in the literature has been that there is a gene – or genes – that controls face and leg markings. The most current study was done in the Franches-Montagne breed, and was published in 2008 (the link will take you to the full article).  After analyzing 23,019 horses, the study found that:

Our data support the segregation of a recessive single gene accounting for 20–80% of the total heritability for the traits under study (head, forelimbs, and hindlimbs markings).

Our association analysis indicated that the putative major gene for white markings is located at or near the KIT locus. However, further studies are necessary to prove that the KIT gene indeed is the putative major gene for white markings. Our association analysis indicated that the putative major gene for white markings is located at or near the KIT locus. However, further studies are necessary to prove that the KIT gene indeed is the putative major gene for white markings.

This and the earlier Woolf studies on Arabians confirm what I have seen over the years while looking for body-spotted sabinos. Like the splash-like horses that never seemed to pan out when it came to what I was looking for (what turned out to be the homozygous SW1 pattern), not all families or breeds with markings pan out when looking for body spots. In fact, some families that consistently produce what most would call sabino face and leg markings only rarely produce body-spotting, and then often just a bit of white on the belly. This phenomenon was noted by Dutch researchers looking at sabino in their breeds, too. In my book, I referred to that particular pattern as “Flashy White Sabino”. I have wondered if whatever causes this type of pattern is simply interacting with (boosting) the marking gene proposed by the team that investigated the Franches-Montagne horses.

But the fact is that while it is commonly accepted among researchers that there is a separate “marking gene”, we do not have a clear picture of the relationship between those markings and patterns. We do have much better tools now, not only with a wider arrange of genetic tests, but also online databases that pair photos and marking diagrams with entries. Just this past week I was combing through one of those resources in hopes of unlocking some mysteries about a breed that consistently has extensive face white without leg white. But the time investment in this kind of research is huge, especially when the pay-off is just clues. Lab work would still be required to get final answers. Until more work is done, though, absolute statements about what is “always” or “never” true should be treated with appropriate skepticism.

Birdcatcher

 

In the previous post, I included an image of a really interesting Greyhound with white dots on his coat. In horses, small white dots on a colored background are often called Birdcatcher Spots. They are not usually as large or as abundant as the spots on that particular dog, which is probably why they go unmentioned in most registry applications. The horse above is a good example of this kind of spotting. You might want to click on the image to get the larger version, and even then the small spots on his hindquarters, barrel and neck are easy to miss if you are not looking for them.

Here is a close-up of some similar spots. Both the horse above and this one are Saddlebreds, where the trait is not uncommon.

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To my knowledge, there have not been formal studies on Birdcatcher Spots. My own experience has been that they are more often seen on chestnuts than any of the black-based colors. (A reader did share an image of a dark bay with very prominent white spots on our Facebook page back in late December.) The spots also seem to occur more often in the “thin-skinned” breeds with finer coats – breeds like Thoroughbreds, Arabians and Saddlebreds. Some have noted that horses with these flecks are more prone to getting white hair growing back over skin abrasions. The horse above did have what appeared to be a few minor abrasions that were growing white hair. A similar tendency might also explain why some roans become covered in dark lines and specks, since injuries on roans tend to grow back with dark hair rather than white.

That does not seem to be the explanation for all cases, however. Here is another Saddlebred from the same show, with unusual white spots on her face. One is just visible on her right ear in this image, and another just behind the browband. A third can be seen in front of the cavesson, and there was another on the left tear bone (not visible here). When asked, her owner said she had always had odd spots like this. Other than her broken blaze, she was a seal brown horse with no other white markings. (She was a very striking horse with a slightly baroque head shape that reminded me of some of the older images of carriage horses I saw while writing my book, and I would have happily taken her home with me!)

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Here is another instance of random white spots, this time on a tobiano.

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Although this fellow had roan hairs around the edges of his pattern, a bit like the horse at the top of this post, and he did have the kind of blaze that might suggest that one of the sabino patterns was present, these were the only round spots like this on his coat so they seemed almost out of place there.

It seems likely that there are multiple causes, either environmental or genetic or some combination, that cause white spots. Reports of how the spots appear, and whether or not they are permanent, vary. Some owners report that their horse was solid colored and then became spotted with a single shedding. Others report that the spots appeared over time. Some say the placement of the spots shift with each shedding. Still others seem to have spots that get progressively larger and more roaned over time. Unfortunately with subtle color variations like these, it is often hard to assemble enough information to draw firm conclusions.

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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.

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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.

KerryBlue

 

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.

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The official Leopard Complex test is now available for $25 from UC Davis. A paper detailing the causative mutation is expected in the near future. Congratulations to Sheila Archer, Dr. Rebecca Bellone and the Appaloosa Project on the discovery!

For more information on Leopard Complex, including some older posts that explain how it works together with patterning genes to produce the variety of appaloosa coat patterns, can be found by clicking on “Appaloosa” under the Categories menu to the right of this page.

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