Considerations on Population Genetics in Dog Breeding
Relating Especially to the Siberian Husky
And the Seppala Case
by Hellmuth Wachtel, Dipl. Ing., Dr. Agr.

THIS 1995 BRIEF WAS SUBMITTED to the Canadian Kennel Club in support of the application to register SHAKAL IZ SOLOVYEV, Russian import Siberian. The author, Hellmuth Wachtel, Dipl. Ing., Dr. Agr., is a well-known personality in the worldwide canine fancy. His writings regularly appeared in “Our Dogs,” “Dog World,” “Victoria Canine Gazette,” “Unsere Hunde” and other internationally known publications. Dr. Wachtel was born in Vienna in 1925, an Austrian citizen and a graduate of Vienna Agricultural University. Retired from his career as a manager in the agricultural industry, he became more active than ever as a voluntary collaborator of the Österreichischer Kynologenverband (Austrian Kennel Club), bearer of the Honorary Pin of that organisation, and collaborator at the Vienna Zoo. His function in the ÖKV was the review of the foreign canine press and the regular contribution of articles on genetics. Dr. Wachtel has lectured on population genetics in Austria, Germany, Denmark and Sweden. He is coauthor of the first Trilingual Pocket Dictionary of Canine Terminology and author of the computer programme “Kyonics” on canine population genetics. Founder of the ÖKV Working Group on Canine Cognition, he was also a canine behaviouralist and consultant. We were very proud to have his contribution appear in SEPPALA NETWORK and were grateful for his positive interest and intervention in the matter of the registration of new Siberia import stock. Dr. Wachtel's work in making dog breeders and national registries more aware of the realities of population genetics has been of the highest order of importance. We are proud to support his efforts.

 

Introduction: Present Situation of Dog-Breeding in General

CONVENTIONAL PEDIGREE DOG BREEDING today centres unilaterally on Mendelian genetics in an individual basis only: the objective is to achieve offspring as near as possible to the breed standard respectively also with high performing abilities in working breeds. This is achieved by:

  • exclusively breeding with registered pedigree dogs (most studbooks are closed, often for a hundred years or more);

  • selecting breeding stock according to show results and scores won in trials (if applicable);

  • extensively using such elite stock, quite especially champion males, in reproduction;

  • practising inbreeding methods (incest, line and back breeding) in order to effectively fix desirable genes.
  • Thus, findings of the relatively recent discipline of population genetics, being in strict contradiction to these conventional methods, are widely ignored or even rejected by breeders, as it is feared that strict implementation of its rules would handicap rapid achievements toward breed improvement as concerns standard conformity. Concepts of "freedom to breed as one likes" also play a part, as well as occasional opposition for commercial reasons, e.g. to lose stud fees if stud use is restricted.

    Unfortunately, as a result:

  • closed stud books have created strictly isolated breed populations which, owing to lack of genetic inflow and small genetically effective population, steadily lose genetic diversity (polymorphism) by genetic drift -- the first case of "inbreeding effect without real inbreeding";

  • extensive use of excellent studs often seriously diminishes the genetic heritage of a breed, shrinking the effective genetic population (this figure never exceeds four times the number of males, so one fictitious male on an indefinite number of females would result in an effective population of just four) -- "genetic genocide," the second case of "inbreeding effect without real inbreeding;"

  • apart from these factors influenced by the breeders, today's breeds experience genetic reductions or bottlenecks -- the third case of "inbreeding effect without real inbreeding" -- since:
    (a) being originated from a small number of founders (a genetic bottleneck),
    (b) they often suffer also from demand- or situation-induced genetic bottlenecks (loss of public appeal, wartime conditions, etc.), and
    (c) intensive selection because of inherited disease control or change or reinterpretation of the standard also can seriously diminish both the real and the effective genetic population.
  • As a result, all conventional purposeful or inevitable breeding conditions entail a constant drain of genetic diversity with every generation. Inbreeding coefficients rise even if no direct inbreeding takes place! Therefore, defect genes of inherited diseases and abnormalities constantly increase; so do lethal, sublethal and subvital genes; thus genetic diseases increase, vitality decreases.

    The situation is quite comparable to that of rare endangered animal species. Regrettably, when multiplying the number of individuals in breed or strain, the effective genetic population increases at a much slower pace than the real population number. So, the effective population of the German Shepherd Dog has been calculated to count today between 400 and 600, in spite of a population of millions of animals, as a result of the above cited factors. Thus, probably every pedigree breed is genetically a rare breed. While the domestic dog possesses an enormous genetic diversity in over 400 breeds, this is genetically ineffective as long as breeds are strictly isolated.

    In contrast, most wild animal populations (unless population and living space conditions are impaired by human activity) normally avoid any close inbreeding by appropriate behaviour (e.g. emigration of one sex of the progeny, mostly males in mammals, females in birds) and assure persistent genetic exchange by emigration and immigration between demes. Without this, the populations would face reduced competing ability and inability to adjust to new environmental challenges, and finally even extinction.

    How International Dogdom is Facing the Problem

    IN MANY COUNTRIES, in conjunction with the advancing animal rights movement, pressure to apply countermeasures is rising. In many advanced countries in Europe it is illegal to breed animals in a way liable to produce offspring with impaired well being, suffering or premature death. Only this spring, the Council of Europe has urgently proposed to ban any "cruel breeding." Already in Germany, a provincial government has prohibited the breeding of merles, hairless dogs and Chihuahuas for that reason. Most significantly, at the recent World Conference of Kennel Clubs held in Ireland, Herr Uwe Fischer, president of the VDH (German Kennel Club), in a speech that raised much attention stressed the absolute necessity to accentuate efforts for breeding healthier dogs, also in order to obviate the menace of governmental actions in this field.

    While action is for the moment still concentrating on abnormalities in breeds and standards, one country (the Netherlands) has already considered a ban on line breeding. The Finnish Kennel Club has introduced regulations of population genetics in its PEVISA control programme for inherited diseases, viz. non-registration of litters with too high an inbreeding coefficient and/or with parents exceeding a breed-specific limit on number of progeny. So far it has not been enacted due to strong resistance of influential breeders. The same holds true for a tentative introduction of mating limits for studs in Sweden; however, an increasing number of breed clubs in Germany and Switzerland have adopted stud use limits. As a further result of the Council of Europe proposals, Norway has pledged in recent press release to take care of maintaining genetic diversity of dog breeds in that country.

    There is no doubt that the number of such measures will increase as there is a growing pressure of public opinion and, in addition, the virulent problem of breeders' guarantees for sale of healthy pups. In Sweden, insurance companies have introduced three rate categories for breeds according to their veterinary expenditure requirements, in order to motivate breeders to care for better health in their dogs.[Ed. Note: Veterinary medical insurance, although practically unknown in North America, is common in European countries.] Besides, "close breeding" is almost becoming a four-letter word in many breeding circles.

    For the first time, in Scandinavia six popular breeds have been DNA tested for genetic diversity; the result is in process of publication. A first doctorate thesis of this kind has been published last year in Switzerland on the Bernese Mountain Dog and the Newfoundland dog, revealing the loss of genetic variety. The microsatellite method of testing was found to be a most useful tool to assess both genetic polymorphism and genetic distance (kinship) of individual dogs to help to achieve mating of least related partners and to find individuals or strains with higher genetic polymorphism, in order to reduce inherited diseases and inbreeding depression. Population genetics is the more in urgent need as expected detection of defect gene markers by DNA tests will inevitably entail a further substantial loss of genetic variety in many breeds.

    Most unfortunately, in contrast to rare zoo and wildlife species, polymorphism tests of canine breeds are still neglected in favour of defect marker detection -- an important objective, but futile without accompanying determination of polymorphism. Genetic screening alone cannot solve health problems without simultaneous measures for boosting polymorphism levels.

    In some breeds, inbreeding levels are so high that outcrossing is not feasible within the breed! As a last resort, some Kennel Clubs are beginning to be less strict against breed crossing. Thus a Newfoundland was used as an outcross for Bernese Mountain Dogs in Switzerland after the war (the Newfoundland being regarded -- wrongly? -- as a close kin to the Mountain Dog), and currently Tervuerens are being crossed with the rare Dutch Shepherd Dog. In both cases, results are satisfactory; a fourth-generation mix in Holland has even been awarded not only Championship but also B.I.G.

    However, inbred animals of the same ancestry do not possess equal levels of homozygosity, which varies according to random influences. Selecting heterozygotes counteracts random drift, so I suspect that highly performing dogs may be more heterozygous than bad performers, since the former are probably heterozygous for a number of vitality genes. (This supposition should be substantiated by DNA research comparing top and poor racing dogs of comparable descent!) Performance selection could help, but not remedy, genetic depletion, at least according to our present knowledge. Introducing one or two non-related dogs per generation into a medium-sized population should help enormously and prove adequate. Highest heterozygosity (and thus heterosis or "hybrid vigour"), however, should result from a three-way cross of three different breeds or very distantly-related strains.

    In the dilemma between absolute breed purity and urgent need to raise the health standard, in cases where pedigree studies or molecular tests do not reveal opportunities for outbreeding within a breed, it will in future become more and more imperative to adopt the option of careful outcrossing with a closely related breed. No doubt the general public is beginning to realise and recognise that pet animals are entitled to an appropriate genetic outfit (genetic diversity), giving them a normal chance to live in good health and to enjoy well being. In this respect and considering the ban on breeding sick or abnormal animals, special breeding laws for pets and/or dogs are called for. These already exist in advanced countries for farm animals, who are at least genetically better cared for than pet animals that are at the mercy of the breeder, the current fashion and the buyers' taste...

    Comments and Conclusions for the Specific Seppala Case

    MR. BRAGG'S BRIEF ["The New Siberia Imports" submitted to the Canadian Kennel Club] reports an alarming genetic depletion for all strains of the Siberian Husky, show type, working type, and the Seppala strain. Crossing of these strains is not desirable for a number of reasons, and besides this would not be very helpful as they appear to be too closely related. Provided DNA tests could not detect sufficiently non-related individuals (there is scarcely a chance), the only way out would be outbreeding with a close relative among another sled or Nordic dog breed, like Malamute, Eskimo dog or perhaps even a phenotypically similar Laika.

    Thus, in the interest of the breed's future, an enhancement of polymorphism is imperative. So it is a lucky chance if aboriginal, autochthonous representatives in tolerable conformity to the standard would still be available. This seems to be the case. The rapid transformation of tribal life in the breed's home region in Russia makes it a unique, short-lived opportunity that could not turn out otherwise than beneficial to the breed. Of course a positive decision should require rigorous follow-up, control and planning of the breeding procedures (back breeding to non-related Seppala, and possibly also non-Seppala individuals).

    In the light of the fact that the animal in question shows good type as well as performance and appears to represent a much more "genuine" Siberian Husky than much of today's "cultivated" strains that have changed in following the reinterpretation of standards usual for nearly all our pedigree breeds during the decades, this chance should not be missed. As a genuine admirer of this marvellous breed, I sincerely hope and trust it will not.