HARLEQUIN BREEDING FORBIDDEN IN GERMANY

a response to the ban by Friedmar Krautwurst,

translated by J.P. Yousha & Monika Olbrisch

with assistance from Katja Hasslebach

(Ed. Note; There is currently a ban on breeding any dog displaying merle or harlequin patterns to another dog of like appearance. This ban is the result of political pressures brought to bear by groups reportedly concerned that the genes involved result in "defective offspring" and thereby, by definition, constitute "cruel and abusive breeding practices". Harlequin to harlequin breeding was banned by the Great Dane Club of Germany (DDC) in the fall of 1995. The following article is a response to this ban by Dr. agg. Friedmar Krautwurst, Dipl. Landwirt, a well established scientist and dog breeder in Germany and author of Die Deutsche Dogge. The article originally appeared in German in DER HUND under the title "Zuchtverbot fuer gefleckte Deutsche Doggen" in September 1995. The following translation was provided by J.P. Yousha and Monika Olbrisch. The translators would like to thank Katja Hasselbach for providing us with the original article and editorial comments. We are grateful to the author for granting permission for publication of this translation.)

I will begin by anticipating my argument: there is no well established genetic knowledge concerning dog breeding to serve as a foundation to the recently enacted breeding ban. This is not to imply insincere intentions to those political and animal welfare activists behind the recent breeding ban; for it is undisputed that the gene producing harlequins and merles, when present in a double dose can result in defects of the sensory system. But what one must certainly condemn is the hasty classification of the breeding of harlequins as "abusive" without specific evidence.

It must be noted that in the scientific literature no direct examination of the Great Dane is contained within the so-called "reports of suffering and damage". From the small number of animals examined, which did not include Great Danes, but rather dappled Dachshunds, it is very risky to extrapolate to a final statement concerning the legislation of dog breeding in general. First it would need to be determined if an extrapolation from the Dachshund population to other breeds is even justified. Do we even know whether the genetic inheritance from breed to breed of identical phenotypic characteristics is identical, totally different, or even results from completely opposite effects?

Knowledge concerning the mode of action of the merle factor when present in a single dose, as exists in the Harlequin Dane, remains unanswered in the current literature, as do questions concerning penetrance and expression. For the accountable representatives of political and animal welfare groups, it suffices that breeding dogs of these coat colors is to be labeled as "abusive". Their actions hence result in the opposite of their intended goal; the destruction rather than the preservation of a type of dog with more that a hundred years of cultural and historical significance. The Harlequin breeder should take this particular opportunity to enlarge upon his genetic knowledge concerning the merle factor.

The merle factor is considered to be a dominant genetic mutation which inhibits, partially or completely, the full pigmentation of the coat. The partial inhibition of pigment appears phenotypically as a harlequin, a blue merle, or other patched or spotted coat types; depending on the base coat color and what other recessive genes are present. If one begins by describing the homozygous black dane and designates this type dog as mm, then designates the harlequin as Mm genotypically, the presence of the dominant M allele results in the partial reduction of pigment. With the complete mutation of both alleles to the merle factor, a genotype of MM exists; a near total loss of pigment and a white Dane resulting. Therefore can one deduce, that in harlequin breeding there are three different gene pairs present producing three different phenotypes:

mm - Blacks = do not carry the merle factor.
Mm - Harlequins = single factored for merle, i.e. heterozygous for merle.
MM - WHITES = double factored for merle, i.e. homozygous for merle

Clear and specific knowledge about the merle factor and its relationship to structural defects has particular significance for the breeder. These defects appear in the form of sensory organ anomalies, such as variable congenital eye and ear defects, as well as reproductive disorders. Because such defects can be problematic, and in that they may not be apparent immediately, the cause and connections of the desirable and undesirable features that result from the merle factor and their mode of operation in both homozygotes and heterozygotes needs to be established. The linkage through which the merle factor mutation produces both loss of pigment and anomalies of the sensory organs can be accounted for by the connection between the shared source of melanocytes and nervous structures in the early embryonic neural plate. The resulting association in individual animals of reduced pigmentation and sensory anomalies can be derived from this early association in the neural plate. Simply stated, these defects can be attributed to the development of early neural structures, which will later develop into the sense organs, in close association with the initial stages of pigment development.

In contrast to the white Dane (i.e. MM), which carries the merle factor in a double dose, the association of the occurrence of sensory anomalies with the breeding of the harlequin (i.e. Mm) Dane, and thereby classification of such breedings as "abusive" cannot be established. The German Great Dane Club (DDC) has determined that, in the future, as a prerequisite for breeding harlequins, a hearing test must be furnished. A similar method for a vision test is currently being prepared. Such testing on a broad basis will provide greater knowledge concerning the genetics/breeding of harlequin Danes. It may be possible to ascertain whether there is an association between the location and size of pigmented areas and sense organ functioning, and whether this is a general phenomenon or varies from individual to individual. It is also possible that differential penetration of the merle factor gene results in varying effects on the structure of the sense organs.

A broad measure taken by the German Great Dane Club (DDC) is the prohibition of breeding harlequin (Mm) Danes to each other to avoid the appearance of white (MM) Danes. The breeder should clearly understand that, statistically, according to Mendelian laws of inheritance, the mating of two harlequins will produce 50% harlequins, 25% white and 25% black Danes. From an analysis done over 11 years and including 2200 progeny of harlequin to harlequin matings, a 14.6% ratio of white Danes was established. These results confirm the assumption that the merle factor in double dosage (i.e. MM) is sub-lethal.  A high mortality rate between conception and birth (i.e. embryonic death) results for MM dogs. The currently imposed restriction to mating harlequin only to blacks also results, on average, in 50% harlequin and 50% black progeny. Double factored white Danes (i.e. MM) cannot result from such matings. Wise selection of breeding stock from these breedings can also produce well marked Danes. Blacks from these breedings are free from the merle factor.

This discussion would not be complete without a comment on the widespread confusion concerning the description and phenotype of so-called white (MM) danes and their associated sense organ anomalies. In the vernacular and the literature MM Danes are generally refereed to as "White Danes". This description is lacking in the same way as "Harlequin" as a term for the spotted or patched Dane. In the German breed standard the "White Dane" is defined as "white Dane without any black markings (albinos)". Both portions of the definition are faulty: a completely white, doubled factored (MM) Dane is rarely found, and albinos do not exist in the breed. In distinction to the above description of merle factored dogs, albinos are produced, if, at the C Locus, both alleles are recessive. In such a case, the chemical reaction that produces tyrosine, a precursor of melanin, the chemical foundation for pigment, is prevented. Therefore any trace of pigment will fail to appear in albinos.

The genetic definition of a double merle factored Dane and its associated phenotype is very complicated and uncertain. I know of animals recorded as White Danes which have no associated sensory defects of any kind. The accompanying photo shows one such individual swimming, who has no equilibrium disturbance due to deafness and possesses normally functioning sensory organs. White dogs, particularly when the result of matings between Boston or piebalds with Harlequins, can also carry the merle factor in a single dosage or be free from the merle factor In the latter case white markings will be determined by other recessive genes. A discussion of the variety of other possible exceptions is not within the scope of this article. If the breeder strictly observes the new provisions of the German Great Dane Club (DDC), there will be no more basis for the classification of the Harlequin Great Dane as "abusive-breeding".

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Copyright 2002 of translation by J P Yousha and Monika Olbrisch, CHROMADANE.  All rights reserved.  Our thanks to the willingness to share this article for educational purposes.