(solid color) and white
“It’s amazing that with all the variations possible, it is only one gene that puts white markings on any solid dog. This gene, as explained by Clarence C. Little, Sc.D. in The Inheritance of Coat Color in Dog, is called the “S” gene. The S gene is a tenacious little critter that carries around the built-in blueprint which he’s not about to change for anyone. The rigidity of the blueprint is best understood by looking at the drawing below. This drawing demonstrates the effect of the S gene on the Black vs the Harlequin. It’s as if the white is spreading over the animals in a predetermined course; and that is exactly what it’s doing.”
“You can visualize the S gene at work (by the above drawing). You can also use it to view the breeding possibilities that supplement each other. It underlines that the black and white spectrum is an inseparable part of the Harlequin genetic pool.”
“Regardless of human will and endeavor, the S gene will continue to appear and work through the Harlequin breed of Great Danes. It’s a matter of intensity. The middleground of C through F is closest to the ideal for both Harlequins and Bostons (Mantles).”
Sounds funny to refer to the bottom Dane on the right of the drawing as a "black" - but that's exactly what he is - albeit his black is "modified" greatly by the "S-gene" - but he is STILL a genetic black.
Just like the "extreme piebald" Dachshund puppy below. Although he appears mostly white (due to the "S-gene"), he is a genetic "Red."
As an aside note, to visualize the "S gene" in Dachshunds in the above drawing of Danes:
– the top dogs of the right hand column are called “Tuxedo Piebalds” or “Irish Spotted Dachshunds”
- the dogs in the middle to lower part of the right hand column are called “Piebalds” or “True Piebalds”
- the dogs at the bottom of the right hand column are called “Extreme Piebalds” (like the one in the photo above)
It's so easy to have a narrow focus of vision when you concentrate solely on a single gene in only one breed. But by examining the gene in two or more breeds, you begin to get a better and more complete overall understanding of how that gene actually works.
The cleanest Harls owe much of their "clean color" (where the white is WHITE) to the S gene (aka piebald gene).
Take a look at Ador Viktoria, his gorgeous clean white color. Then look at Prinz Fuchs, Ador v Amalienborg (and others) and you can SEE the S gene at work in those gorgeous Danes. You can even trace the S gene back through the progenators of those dogs, simply by studying their photos.
Take a look at the incredible Dane "Ador Viktoria" below:
This dog was not a harl at all, he was a mantle (you can tell by the facial markings), and represented the "S gene" at work in the above drawing (the mantle dog on the right labeled "F"). On the "Danes of Special Note" you can see on his page the incredibly "clean colored" harls he produced. His fantastic lineage was spread worldwide to the best Dane breeders of the time and thankfully has been passed down over the generations.
Where did he get at least one "S gene"? Well, by studying photos of his progenitors, it seems fairly obvious that his grandfather "Prinz Fuchs" was a likely candidate:
And, as a matter of fact, Prinz Fuchs was the identical version of the "S gene" factor of Ador Viktoria but in a HARL (versus a mantle). Take a look at the "F" harl in the above drawing.
In Great Danes, the "S gene" is an integral component of Mantles - and of clean-colored Harls. To brag about trying to "eradicate it" is horrifying.
I am utterly dismayed by some of the emails I get from Dane breeders who seem on a crusade to "eliminate" this age-old and completely natural "S gene" from their dogs, and obviously without understanding the contribution that it makes to Harls and Mantles.
It's just one of a number of genes which have been identified and which "purists" seem on a mission to obliterate.
How very sad....and how very misguided.
There is a lot of hullabaloo surrounding them. But guess what? They were most likely NOT caused by any "natural" mutation - they were the results of the domestication process by man.
Take a look at the following link, an experimental study of domesticating foxes was done to explain the physiological and emotional differences between "wild canines" and "domesticated canines." Besides the obvious decrease of sexual dimorphism (the tamer they became by generation the more they LOST their distinctive male vs female body size and skull shape), the tails became shorter and curlier, etc. And one of the FIRST differences noted, was an increase in "depigmentation" (ie more white, less color). WHY?
"Belyaev determined that this piebald pattern is governed by a gene that he named Star. Later my colleague Lyudmila Prasolova and I discovered that the Star gene affects the migration rate of melanoblasts, the embryonic precursors of the pigment cells (melanocytes) that give color to an animal’s fur. Melanocytes form in the embryonic fox’s neural crest and later move to various parts of the embryo’s epidermis. Normally this migration starts around days 28 to 31 of the embryo’s development. In foxes that carry even a single copy of the Star gene, however, melanoblasts pass into the potentially depigmented areas of the epidermis two days later, on average. That delay may lead to the death of the tardy melanoblasts, thus altering the pigmentation in ways that give rise to the distinctive Star pattern."
So what changed the migration rate of the melanoblasts to begin with?
Answer = Reproduction.
Wild dogs bred once a year. The whole purpose of "domestication" is to increase reproduction, and the captive foxes (and early domesticated dogs) became sexually mature earlier and also became more fertile - allowing 2 seasons a year instead of one. This change of increased reproduction changed the rate of migration for the melanoblasts and created genes for piebald and "mottling" (aka dappling or merling - and remember, a "harl gene" is an offshoot of the "merle gene").
The article has fascinating photographs of the differences of coat color (ie the increasing areas of white and also "mottling") over the generations of foxes tested. (ever see a piebald fox? you will there!) And it must be noted that they were not inbreeding the generations over and over. They started out with 40 male foxes and 100 vixens, and by the 8th to 10th generation, noted the distinctive changes in the coat color.
Anyway, it's a fascinating study, for anyone wanting to pursue it. And reinforces the affirmation that piebald (or "s-gene" as it's now called) and merle/dapple (and hence harl) were patterns which were developed during the early domestication process of the dog (and likely within the first 8 - 10 generations of the earliest domestication).
They were NOT unexplained "mutations"
They were NOT intentionally "created" by man - though his disruption of the canine's normal reproduction cycle caused their creation
They are NOT a result of inbreeding, poor breeding or poor dogs.
They ARE an inherent result of the domestication process, dating back to when the dogs were FIRST beginning to be domesticated, and their breeding controlled by man.
All the old-time harl breeders KNEW that merle could exist without harl - but not vice-versa. They didn't have the "more book learning than sense" that the younger breeders nowadays have...the old-time breeders knew because they EXPERIMENTED with their breedings.
But this goes back to the difference between colors and patterns - you HAVE to know the difference in order to understand the genetics. Anyone can spout off the ee or bb or DD genetic codes...but most of those people don't understand HOW they work OR the difference between "colors" and "patterns."
And if you don't know HOW the pattern genes were created, and HOW they work, and HOW they are different than color genes - then you can NOT understand genetics.....period
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