101 Wolfdogs   

(Part I - History, Literature Search, Coat Color Results)

by Ann Dresselhaus
Reprinted with permission of the author.

Review of:  Journal of Genetics, Vol 42, No. 3, pages 359-427, 1941,
Wolf-Dog Genetics,  Prof. Dr. N. A. Iljin

From the same country that brought us the 40 year Fox Domestication Project comes an 8 year venture into the genetical analysis of 101 wolfdogs.

In 1923,  about thirty years before the Russian scientist,  Dmitry  K. Belyaev was to begin his project  which would show that wild fox could be domesticated in as little as 40 years, another Russian scientist,  Dr. N.A. Iljin who was the Director of the Institute of General Biology at the First Institute of Medicine in Moscow began his 8 year breeding experiment to facilitate the collection of data on the genetics and morphology of wolfdogs.

Improving local dog races:

Dr. Iljin did a literature search before he began breeding and found quite an impressive record of wolves being crossed with dogs down thru time beginning as far back as Aristotle in the fourth century B.C.   Pliny in A.D. 23 - 79 reported that the Gauls attached their bitches to trees in order to have them mated with wolves. These practices continued into the 18th and 19th centuries.  Iljin found that the breedings described were not investigated in a scientific way but were either occasional experiments or were directed toward the improvement of local dog races including the Eskimo dog, the Indian dog, and the Hungarian dog.

Many wolf-dog crossings were carried out in zoological gardens and parks such as the Moscow Zoo Park, Jardin des Plantes in Paris, Hanover, Stockholm, Halle, Brehm, and the Marseilles Zoological Gardens,  where they were regarded in a "well-respected manner" according to Iljin.

Author's note:

The German Shepherd Dog Mores Plieningen, SZ #159, who was bred to the first Stud dog, Horand Von Grafath (previously known as Hektor Liksrhein) and whose blood is said to be in the pedigree of every German Shepherd Dog in the world today, was the granddaughter of a wolf at the Stuttgart Zoological Gardens.  Their son, Hektor Von Schwaben, SZ #13, figured heavily in the early German Shepherd Dog line.

The beginning:

A wild caught 'zonar-grey' (black phase with grey highlights) male wolf was mated to a black female sheepdog mix resulting in an F1 generation of 13 puppies.  Half of the F1 generation had the mother's  solid black coat color or were black and tan and half had the father's zonar-grey coat color.   There were 61 puppies in the  F2 generation, 24 in the F3, and 3 in the F4  for a total of 101.  The coat color of 73 specimens was exactly

Mapping Iljin's findings to modern day canine coat color series:

It appears that what Iljin referred to as the 'zonar-grey' coat color is the expression of the same allele that Burns and Fraser (1966) refer to as the 'ag' agouti (wolf grey) allele in the Agouti (A) series.  The black and tans that appeared in Iljin's F1s were either expressions of the 'as' (saddle marking) alleles or the 'at' (bicolor) alleles in the same Agouti series.  The leading (most dominant) member in the Agouti series is the A (dominant black) allele and the third most dominant member allele is the 'ag' agouti (wolf grey) allele.   However, interestingly enough it appears that the solid black color that the zonar-grey wolf male carried as a recessive and the matching allele in the black female sheepdog are indeed evidence of *another*  black  allele (recessive) 'a' in the same series. The 'ag' agouti (wolf grey) allele is dominant to the 'a' black (recessive) allele.    A good reference to get a handle on these coat color series is, "Genetics of the Dog"(1989) by Malcolm Willis which on  p. 65 explains the Agouti (A)  Series.

Recessive Black allele found in wolves, German Shepherds, and Belgian Shepherds:

If black were dominant then all black dogs would have at least one black parent.   Willis (1976) showed that only 55 in 115 black German Shepherds had at least one black parent.   In Iljin's experiment, 2 black F1 hybrids (apparently a/a) produced only black offspring, while 2 F1 zonar-grey hybrids (apparently ag/a) produced both zonar-grey (ag/a or ag/ag) and black (a/a) offspring which is indeed evidence of the recessive black
allele 'a' in the wolf.

Carver's (1984) data show clear evidence of a recessive black allele in the A series using data from 564 black German Shepherds, which in addition to the Belgian Shepherds  are 2 of the very few dog breeds known to carry the recessive black allele.  This may be considered circumstantial evidence of the influx of wolf heritage into the early dog lines of the Belgian Shepherd and the German Shepherd.

Doberman markings and liver-colored wolfdogs:

Iljin found the tan markings in the F1 offspring (probably at/at) to be identical to that of the Doberman (at/at) and the Gordon Setter (at/at) which is the expression of the 'at' (bicolor) allele in the same Agouti series.   These markings were 'covered by' the recessive black 'a' allele which makes it dominant to the 'at' (bicolor) allele.

Iljin describes 'brown-fulvous' colored F2s with light whitish-blue irises that were the progeny of 2 black (F1) wolfdogs.  The manifestation of this color was independent of other characters indicating an independent allelomorphic site/series. It is not clear to me whether this is the expression of what is in the present day referred to as the liver/chocolate allele 'b' in the B series (Little 1914), the Dilution allele 'd' in the D series (Little and Jones 1919), or the chinchilla allele 'ch' in the C (Albino) series (Burns and Fraser 1966).  The definition of fulvous  is "yellowish-brown, tawny, dull yellow, or yellow";  however, the black and white pictures of this cross looked quite dark - nearly as dark as the black coat color and the nose color could not be determined from the

This 'brown-fulvous' color would be confirmed to be an expression of the B series if Iljin would have noted that the nose color of his 'brown-fulvous' offspring was not black but brown or liver; however,  there was no mention of nose color at all.  He did mention that the coat color was similar to Newfoundlands, Pointers, and Dashhunds, all of which can occur in the liver/chocolate color phase indicative of the B series allelle 'b'.

Whitish-blue-eyed wolfdogs:

Iljin mentioned that the eye color of all these 'brown-fulvous' wolfdogs was similar to the eye color of the 'marbled' Great Dane, which I think is what  we call the Harlequin Great Dane today. However, no Great Dane today (Harlequin or otherwise) is *supposed* to carry the 'b' allele, so the light eyes are likely the effect of the Dilution or D series.  The 'd' dilution allele is recessive to the 'D' (regular intensity) allele and often lightens the eyes as well as the coat which could ALSO explain the light eye and ('brown-fulvous') coat color of these F2 offspring.

The last possibility is that the base coat color of the offspring in question was lightened by the 'ch' chinchilla allele (in the C series) which does not allow the full expression of coat and eye color to take place.  Canines whose coat is lightened by the 'ch' allele which is recessive to the C allele (full expression of coat color) can still have black noses.  There is also a postulated allele in this series 'cb' cornaz coat/blue-eye allele (Pearson and Usher 1929) which can give rise to blue eyes and is recessive to 'ch' chinchilla allele.

'Brown-fulvous' coat color, whitish-blue eyes, spots, and wavy coats are likely dog contributions:

It should be noted that regardless of the cause of the lighter ('brown-fulvous') coat and eye color the source of the recessive allele(s) does NOT have to be the wolf since 2 F1s were bred together to produce these lighter coat and eye colors.  Each F1 could have inherited the recessive 'brown-fulvous' source allele from their dog mother and when bred
together this recessive allele was expressed (ie. doubled up) in their offspring.

In much the same way as the 'brown-fulvous' coat color was produced, white-spotted offspring  and longer wavy-coated F2 offspring appeared. When 2 black F1s were bred together, some F2 offspring with white spots occurred although there were no occurrences of white spots in the F1 generation.   This points to the expression of the 'si' (Irish Spotting) allele in the Spotting (S) series (Warren 1927 and Little 1957) which was
probably inherited from the dog mother.  This allele is recessive to the 'S' (Self) allele.  Most wolves are likely S/S although there are sometimes small white chest and feet marks in wolves which could be caused by either the 'si' allele or modifiers at other gene loci that act on the S loci. There is also heavier spotting in the S series through the 'sp' (Piebald) and the 'sw' (Extreme White Piebald) alleles which are virtually never seen in the wolf.

When 2 straight regular-coated F1s were bred together, some F2 offspring with longer wavy-coats occurred.  This is consistent with the modern description of long (wavy) coats as recessive to straight regular coats. Again there were no occurrences of long (wavy) coats in the F1s so the long (wavy) coated allele likely came from the dog mother alone as wavy coats are virtually never seen in wolves.

'Blue' wild wolves:

Ashen 'blue' wolves were mentioned as being caught (but not bred by Iljin) in the northern region of Tobolsk in the USSR.  The were described as being a dilute zonar black or a zonar black with reduced black intensity.   It is my guess that this is actually the expression of the  'd'  dilution allele in the D series discussed above which may occasionally occur in certain geographic regions or among certain subspecies of wolf.  Since it is a recessive allele, it would be nearly impossible to eliminate from the wild
population of wolves unless a single dose of it conferred some maladaptation upon its carriers as well as upon those that expressed it.

The segregation of coat color in wolfdogs proceeds in the same way as in dogs:

Characters recessive in dogs are recessive in wolfdogs.   Characters dominant in dogs are dominant in wolfdogs.  The genes of dogs introduced into crosses behave in just the same way as when they meet their normal allelomorph.  Coat color in wolves and dogs result from identical genes in the same linkage groups.

End Part I  (History, Literature Search, Coat Color Results)
See Part II  (External Characteristics Results)



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