SABLES: GENETICS AND MYTHS
Some time ago, a GSD breeder-judge in Germany wrote an article that was carried on the Internet, entitled “The Sable Shepherd… A Museum Piece?” (liberal translation). Several people have asked me about some of the statements in it, and I thought a magazine or website piece might be the easiest way to answer.
The color pattern we in America call “sable” is called “grau” (gray) in Germany. Neither word is fully accurate, but experienced dog people know what is meant. The strict translation of what is possibly originally Latin, then Russian-Scandinavian in origin, but is found in variations in many languages including “sable” in English and French, is “black”. Actually, even before that meaning, it was the name of a small glossy-black weasel or ermine found in far-northern Europe and Asia. From the Latin zobola, Russian soboli, Scandinavian sabel, and German zobel, we get our word, but not our meaning. It brings a different image in other breeds, such as the Collie, Basenji, and Sheltie where it refers to a reddish-yellow dog. Even in the Pembroke Welsh Corgi, the sable dog gives mostly an orange impression. In the Shar-Pei, we see a color that looks like the GSD sable, but has a different genetic constitution. There is even the white GSD, some of which appear to be genetically modified sables, but certainly without any black.
Let’s define a couple more terms before proceeding. Everyone knows what is usually meant by the word “color”: the reflection or refraction of certain wavelengths of light that tell us the apple is green, the traffic light is red, or the rainbow has seven more or less different hues. But “color” is often the term used when GSD people talk about black-and-tans, blacks, bi-colored dogs, or sables. For that meaning we really should use the term “color pattern,” because some of these dogs may have more than one color or hue in the coat: shades of red/yellow/brown, black, white, and “dilutes” (blue and liver). The German word for what we call “tan” is “gelbe” (yellow). While “braun” (what Americans often braggingly like to call “red”) is also seen in the Ahnentafeln (SV registration-pedigree), it is genetically and chemically the same as yellow or red, altered with hue and intensity modifier genes.
A hair bulb can manufacture more than one type of melanin, and can alternate production in such a way that some hairs, such as most of those on a sable dog, are dark-tipped, followed by a lighter midpiece and an even lighter base (or perhaps a dark base). There may be two shades of yellow: one reddish and the other cream, on one shaft. Sometimes the phaeomelanin (this form produces the non-dark pigment) is concentrated in the tip, and the eumelanin (which produces the black, blue, or liver) is in the base, though not often. The banding or alternating between dark and light sections results in a beautiful variety of colorations in the breed, especially around the neck, withers, and shoulders. The definition of a sable pattern should refer to the guard hairs being tipped with black. The more hairs and the longer that tipping, the darker the dog. The richer the phaeomelanin-influenced parts (undercoat and lower/ventral/leg portions), the more the dog is called a “red sable” instead of a “gray sable” (or faded). The “black sables” are those with much tipping but not always much red underneath.
Agouti is a term borrowed from genetics studies on that type of rat. It refers to the banded-hair coloration or pattern seen in Elkhounds, wolves, and sable German Shepherds, but can also be seen in varying amounts in the neck, shoulder, tail, croup and border markings in saddle?marked dogs. Some saddle-marked B&T GSDs have so much grizzling in the black blanket or so much undercoat showing, that they are hard to distinguish from sables, and some sables that carry the partially-hidden B&T-saddle pattern recessive have so much of that recessive showing through that they look like washed-out B&Ts. At one chromosome’s locus A, the German Shepherd Dog has a pair of genes, one on each of the chromosome pair, which together determine the major markings we call “pattern.” This pair may have two of the same or two different alleles, one more dominant and the other recessive to it.
Ay, sable — The superscript y is used in genetic parlance to refer to the yellow pigment that can dominate the appearance of the tan?sable dog, even though a true sable should have those hairs tipped with black. That black appearance has been lost in many breeds through selective breeding. In other breeds, where a more dominant allele for self-color (solid) exists, the sable or yellow pattern is ay (lower-case a, because the capital letter refers to the most dominant allele in the series). For example, the black Lab is As while the yellow Lab is ay. The color lying under the black tips on a GSD’s coat, for example, is due to other genes. A sable is generally held to mean a dog with black-tipped guard hairs, this banding being obvious over most of the dog’s surface, especially the dorsal parts. In those other breeds I mentioned, the word is used variously to refer to a yellow dog regardless of the presence of any black tipping. Many or most “sable” Collies and Corgis have lost the genes calling for expression of black in the hair. There is also a “white sable,” but that subject is treated in a different article.
as, saddle — The saddle pattern gene is below Ay in order of dominance. The use of lower case shows it is not as strong in its influence over pattern as the sable gene, but we know it is dominant over other alleles in the series. The saddle pattern has been the most popular in the past 50 years. Whether, as the Germans describe it, the base color is black and the tan portions are the markings, or whether (as some of us think of it) the base color is brown/tan and the saddle, muzzle, and head markings are the areas of added pigmentation, it's moot, like arguing about the color of a zebra.
at , bicolor — Again, this pattern, resembling the coloration of a B&T Coonhound or Doberman Pinscher, is best discussed in a separate article. Some think it belongs on a different chromosome locus.
Incomplete dominance — In the A series, incomplete dominance of one A series allele over a lower one is often, though not always, visible to the experienced eye. A heterozygous Ay as sable (i.e., the genes in the pattern pair not identical, and we would say he carries a recessive for saddle pattern) will have a shadow of a saddle with slightly more dark?tipped hairs (or longer black band on those hairs) than the homozygous Ay Ay sable has in the areas commonly marked black in the saddle dog. If the sable has a hypostatic (recessive) gene (at the E locus?) for solid black, such a dog is usually considerably darker in overall appearance than are his lighter gray or golden sable relatives. Karilea's Cito, Dipadon's Dasher, and other sables that carried the black factor and were capable of producing solid blacks had this dark look. Many “working-lines” German Shepherd Dogs carry the black or bi-color recessive (or both?) and have a very dark appearance. It may well be that most extensively-covered very dark sables carry the bicolor recessive, and/or the black factor at the E locus as well. This may be especially true in the working lines, where these three patterns have not been pushed out of the picture by the saddle-marked dogs as has happened in the breed show ring. For greater detail on genetics of coat color and other factors, get a copy of “The Total GSD” at <www.Hoflin.com> or from your favorite book dealer.
Since beginners are often confused as to what will happen if they breed a certain bitch to a certain dog, let’s put some dogs together on paper and see what you get. As an example, the homozygous sable, with both A series alleles (genes in the pair) being Ay, is bred to the homozygous B&T saddle, as as. Regardless of other factors such as depth or distribution of pigment, the Ay Ay X as as (sable-saddle cross) will give a litter of 100 percent sable phenotype, though possibly not as distinctly patterned as the one parent's, and all the pups will carry the as as a recessive. Very few homozygous sable GSDs seem to be running around, so most of the ones you see will produce some B&T pups, even when bred to another heterozygous sable. The variations in shades and markings are influenced by “modifier genes” such as E, Em, e, or eb which govern the extension of black pigment to a mask, and influence on what percentage of the body that pigment will be used. E alleles appear to act more as “advisors” to the A series. There was a nice article on masks and patterns, by the “sable Linda Shaw (blonde, not black-tipped)”, in the July/August 2000 Schutzhund-USA magazine issue.
The Concentration Series, C Locus
C alleles (genes at a location we’ll call “C”) control the color or Concentration of the tan parts of the Shepherd Dog. Newborn B&T whelps often appear all?black until close examination reveals some silvery or tan coloring on lower legs, cheeks, eyebrows, and around the anus, often only at the vent in the case of the darker pups. The tan areas will gradually increase at the expense of the black saddle and other markings, with the process tapering off and nearly stopping at maturity. As the pup grows, the tan creeps up the forelegs to meet the widening chest and neck lighter markings, and up from the hindlegs to the haunches. Eventually, the pup that had a tiny bit of tan on brows, cheeks, and feet becomes a much lighter dog with a saddle halfway down the ribs and just barely covering the hip bones.
Bicolor dogs or saddle dogs with dark recessive modifiers do not usually change as much. Sable puppies generally start off a little lighter at birth than the shade they later develop, but much influence comes from genes at the E and C locations on the chromosomes, and the delayed development of topcoat. “Hypostatic” is like “recessive” but not necessarily on the same chromosome or locus. A sable with a black hypostatic gene (eb) or a bicolor recessive (at) or one of each may be darker at birth than a littermate without such modifiers, other factors being equal, and remain dark throughout his lifetime. A strange paradox has been reported: Dr. Michael Fox has observed that wolves start out darker at birth and get lighter for a while. Many geneticists think that wolves are probably true agoutis, whereas the GSD has probably lost that allele and has the sable gene as its most dominant one. However, I have seen wolf-GSD crosses that are almost indistinguishable from “regular” sable GSDs. The C locus governs the color of the “tan parts,” as I said, whether the dog is sable or B&T. The amount of black coverage in the saddle area is determined by other factors.
Black and tan (saddle or bicolor) German Shepherd Dogs with somewhat faded points or considerably lighter hair on the inside of the legs may have, at the C locus, a “chinchilla” (fading effect) recessive making itself partially visible. The lighter those portions are, the more likely is the presence of such an allele, or even two of them (homozygous for chinchilla). In true sables, cchch would tend to give a very pale gray, light golden, or sooty cream appearance to the hair comprising the “tan points” or that portion of the back’s (blanket’s) guard hairs that is closer to the skin than the black tips are, the depth of color depending on interactions between other genes. The e allele is also involved in the pale solid-golden mislabeled “sables” reported by a number of observers, these dogs being actually B&T in genotype and born with a black and yellow?gold?tan as or even at constitution and appearance, yet losing the black by adulthood. Breeding to a solid black would not necessarily cover the effect of the chinchilla gene in the progeny, for this gene does not act on black pigment, only on phaeomelanin—that is, tan hair. A solid black dog could easily be cchch cchch in genetic constitution with no way of your knowing except for much progeny testing. Because a solid black has no tan showing, there is no way of knowing by looking at it whether its C locus alleles are chinchilla or call for “normal” hue when allowed. A black dog with such paling genes at the C loci would not contribute color depth to tan parts of B&T offspring when mated with a saddle-marked dog, for instance. The same would be true if bred to a sable—if you don’t get strength of pigment at the C locus from both sides of the family, you could have poorly pigmented pups, even if one parent is a solid black. Also, the common claim that breeding to a sable every few generations will ensure good pigment is unscientific; keep in mind that some of the sables you might choose to use may themselves carry the chinchilla gene. So, generally speaking, if you have an otherwise excellent animal with a lack of strong pigmentation in the “tan areas,” it would be advisable to breed it only to dogs of good pigment which you feel do not carry the chinchilla recessive themselves. Otherwise, there may be very little color difference between your stock and the “bargains” found in the classified section of the newspaper.
When the GSD breed got started, there were many sables, probably far more than there were B&Ts, and perhaps some with wolf-agouti pattern as well. (In my book, you will even see a blue merle GSD from the 1920s!) What happened to the sable in the show lines, and why have the “working lines” kept so many of them? The answer to the second part is simpler, and it involves the breeder Alfred Hahn of BuseckerSchloß fame. When I visited with him, he told me how he started his kennel in 1925, and how he had a special leaning toward the “gray” (sable) dog. He showed me many photos of his past and current successes, and in most cases they were sables. Hahn had the single greatest influence on both the preservation of the sable, and on the furtherance of the “total” show-working dog of any in his long tenure. He blended the best working dog lines with the best “show” dogs. Behind the greatest “golden middle” dogs that ever lived (the Lierberg B, D, and other litters) was the BuseckerSchloß name. I knew Bodo, the hardest and most imposing GSD I’ve ever come across, and I had a Gin v. Lierberg daughter that could also eat iron and spit out ten-penny nails. They were B&Ts, but they had tough and beautiful sables in their backgrounds.
Most sables will start out lighter than they will later become, though there are some that show the reverse progression. It may take three years, according to van Dorssen, for a sable to develop its adult color. I’ll never forget the poor air-headed “breeder” years ago in one GSD club who put down most of her litter at birth because she thought she had “those dreaded white dogs.” She had no idea that the pale newborns would most likely grow up to have the same sable coloration that the sire had!
Breeders have noticed in both sables and B&Ts, that continued breeding within the one pattern or the other (such as sable-to-sable or B&T to B&T) will eventually produce many with faded pigment. This is where the misconception arises that you should breed to a sable every so many generations to enrich color. Nonsense! Fading happens in sable-sable and sable-B&T breeding, too. The real situation is that in either camp, breeders were not careful enough to re-introduce intense pigmentation, and as you have heard in other instances, if you don’t use it, you lose it. A sable will not automatically improve depth of color; in fact, if you breed to the wrong sable (or any color pattern) you can get faded pigment in your pups. However, to claim as did that German breeder, “…if, after several generations of purely sable-to-sable cross-breedings, a [B&T] combination is made, no loss of pigmentation will occur,” it ain’t necessarily so. The B&T could be less than well-pigmented, and if he didn’t have better genes for producing deep pigment, especially in the phaeomelanin-influenced coat areas, you could still be taking a step backwards. Nor is that person correct when she agrees with those who surmise “that a well-pigmented B&T who has one sable parent, produces a better pigmentation” for both B&T and sable offspring. Having a sable ancestor, no matter how close, has nothing to do with it. If the dog in question is a B&T, it has NO sable in it… sable cannot be recessive to B&T, and therefore is not present to wield any influence. Those who believe it does, make no more sense than the homeopath who thinks that the “memory” of some molecules that are no longer in a bottle can influence the future contents of that vessel. To say that “The sable gene cannot lose intensity” by continuing to breed sable to sable, is totally unfounded. When one attributes good pigment in a particular dog’s pups “to his sable ancestry on the maternal side,” is likewise ‘way off the mark. The dog’s well-pigmented offspring inherit “modifier genes” for richness of pigment, even some from light-colored females he may have been bred to. Improving pigment “by using a sable female or one of her black-&-brown descendants” is no guarantee, not even an indication of the direction to take. To use as an example a dog that almost everyone is familiar with, the offspring of Timo Berrekasten, that spectacular sable VA dog at the Nuremberg, Karlsruhe, Bremen, and Düsseldorf Sieger Shows, generally appear to have outstanding pigment because of much black overlay, though some could have darker color below the black portion of the guard hairs. However, it is not because he is a sable, but simply because he has such good pigment in much area. Look at his legs, and you can see the richness that is not covered up by the extensive black portions. When I first wrote this for a magazine, I said, “I would love to have a dog like that!” Then I got one: a beautiful Timo daughter who looked and acted like her sire.
Another myth that German breeder stated long ago is that “The dog that dominantly produces sable [homozygous]… is [also] homozygous for short hair.” People in the U.S. have echoed this totally groundless misconception, too, that such a sable cannot produce long-coats. There are sufficient examples that refute this; after all, science (of genetics or anything else) is based on observation and repeated experiments. There is no chemical or genetic connection between length of hair and pattern. They are on completely separate loci. You can almost as readily produce a long-coated white, sable, or solid black as you can a bicolor or saddle-marked dog. I say “almost” only because there are fewer of them around, and fewer cases of people breeding for those expressed purposes. But it is just as easy, if you locate and breed the “right” pair, as it would be to produce a solid-liver long-coat, a blue-and-tan long-coat, or any other such combination of odd coats. To say that a homozygous sable would offer “a way to at least restrict the problem with long coats” simply is not true. The Shiloh Shepherd subfamily of GSDs has many scores of examples of long-coated sables.
Another misconception is that in breeding a homozygous (pure) sable to a B&T, “50% of [the pups on average] would be homozygous for sable, and 50% heterozygous.” Of course, you know that 100% of the pups would be heterozygous sables; i.e., they would all have the sable pattern phenotype, but each will carry one of the B&T’s recessive genes. None could be homozygous for either B&T or sable. The additional claim that “sable dogs are more resistant to strain [sic] than black & brown dogs” is unfounded (She probably meant “stress,” but either is equally unscientific and neither is a statistically valid statement.). There is no connection between color pattern (sable vs. B&T) and working qualities. There is only coincidence—the fact that the same people who had produced sables are largely the same ones who are more into the working-sport activities in the breed. It is concurrent, not cause-and-effect. People with high-scoring sables tend to look for more sables to buy or breed.
I had a problem with some of the numerous false claims and comments made in the “paper” that had been circulated on the Internet some years ago regarding “Peculiarities seen with sable pups”:
“The weight at birth is usually …higher.” Where is the data? I’m not ready to accept that without proof and reason.
- “Clearly higher vitality.” I have not seen that, nor can imagine any logical reason.
- “Lower mortality during the first ten days.” Nor have I seen that. I suspect the assumption is based on coincidence.
- “Higher weight when tattooed.” Not among American dogs at that age, so why should it be so in Germany? No basis in science at all.
- “Less heat sensitivity…” (pups, adults, shows, trials) Maybe less black to absorb heat? Again, poor correlation; pointing the finger at the wrong cause.
- “I have never observed [light nails].” Well, I certainly have! I’ve judged and examined untold thousands of GSDs.
- “More often black spots on or below the tongue.” No difference; this is inherited separately.
- “Never [light] lips or gums. (or) White hair in the ears.” Purely incidental. No connection with coat color pattern.
- “Almost all sable females and many black-&-browns stemming from sables feed their pups out of their own stomach.” Coincidence.
If you, reader, have also entertained thoughts along those lines, I challenge you to apply reason and reliable data to such ideas. Turn the flashlight of science and logic onto superstition and watch it disappear.
In summary, there are many misconceptions about coat color or pattern, and the supposed effects on other characteristics. The sable GSD was making a comeback in the show circles, thanks to SV & WUSV boards of directors and a recent but now-deceased SV president. It has been idolized by many Schutzhund-sport-only folks and the object of adverse prejudice by many show-only fanciers, neither side having any basis in common sense or science. A good sable can contribute much to the breed, especially in the area of genetic diversity. While a GSD does not run on its hair, or bite with its pigment, good color and attractive markings help to give the breed its highly important expression and overall impression. Thus, they contribute to the all-important preservation and promotion of “Type.”
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