Genetic Load
The difference between the fittest genotype of a population and the average fitness of that population is known as genetic load. [Muller, 1950] It is, of course, caused by the presence of lethal, sublethal and subvital alleles. The more such alleles found in a population, the greater the genetic load. Genetic load is sometimes measured by the number of lethal equivalents, and the severity of inbreeding depression can be quantified in this way. Humans in general normally carry in a heterozygous state from 5 to 8 lethal equivalents per person - genes or combinations of genes any one of which, if homozygous, would cause the death of the organism. It should be emphasised that genetic load is present in every population, since never are all individuals maximally fit. The presence of lethal, sublethal and subvital genes is a normal state of affairs in all species. Homozygotes for such genes are usually so infrequent as to have little effect on species fitness. It is only when founder events and inbreeding occur that the gene frequency of deleterious alleles rises and genetic defects start to become a problem as the growing genetic load degrades the fitness of the inbred, limited population. Thus in the case of purebred dogs the problem does not inhere in the presence of “defect” genes, but in the registry and breeding practices of the purebred dog fancy!
Balanced Heterozygous Population Structure
In recent decades growing evidence from DNA studies of protein polymorphism conclusively disproved the “classical” view of species as being homozygous at most loci, with the phenotypes of all individuals of a species conforming to that of a type specimen. Population geneticists and evolutionary biologists now realise that typological concepts are useless in a natural world in which populations may best be described genetically not as individuals conforming to a type but as arrays of genetic variability. Some of the implications of the “balance view” are elucidated by one geneticist as follows:
Species that are diploid and cross-fertilised [this includes all mammals]… characteristically carry large stores of genetic variability in a balanced state in their populations… Genetic recombination naturally generates diverse genetic types from the large field of variability in the gene pool. In order to meet environmental challenges, natural selection in many such organisms tends to develop a system based on the higher fitness of heterozygotes. These are maintained under regimes of selection that exploit the advantages of heterozygosity for many alleles simultaneously. In these, the large amount of genetic variability is continually being recombined as balanced hybrid vigour is maximised…
The genetic system is not a fixed and frozen entity but is dynamic and variable… By its very nature, this genetic system is inimical to the perpetuation of sameness. At each reproductive event an enormous field of genetic variability is produced. Most of the variability is held in sexual populations by a complex balancing selection based on the superiority of fitness of heterozygotes…
The biological conserver, short of putting the DNA into liquid nitrogen, cannot hope to freeze the characteristics of any natural population, be it a deme (local population), a subspecies, or a species.
Hampton L. Carson. The Genetics of the Founder Effect, 1983
Efforts at artificial selection and breeding which attempt to defy this system of balanced heterozygosity and variability will almost certainly fall foul of the kind of difficulties we are now encountering in purebred dog breeds. It is hopeless to attempt to freeze the genetic characteristics of small populations and even the attempt, which is doomed to eventual failure, is quite costly in terms of the loss of hardiness and viability. Artificially selected populations, too, can and should be maintained in a state of dynamic heterozygous balance. Thus the entire problem of genetic defects would be minimised.
Assortative Mating
Assortative mating is a method of selective breeding capable of creating homozygosity for desired traits without having as great an effect on overall homozygosity as does inbreeding. It consists of mating phenotypically similar individuals that are not closely related. This method of selective breeding would be capable of maintaining a reasonable range of breed type in a balanced-heterozygosity breed system with an open studbook.
Having now acquired a few of the more crucial concepts of population genetics, we are prepared to examine in a new light the nineteenth-century system of dog breeding and registration which we have inherited. As we prepare to enter the twenty-first century, perhaps we can conceive a renewed system which will serve our dogs and their breeders far better than the present one.
The Crux of the Problem
As we face the millennium, the one problem which most concerns the entire purebred dog fancy is genetic defects. Breeders used to worry about overshot/undershot bite and cryptorchidism. Not much else of a genetic nature was cause for concern; fanciers were a lot more worried about distemper, hepatitis and internal parasites. Breeding programmes concentrated on individuals’ visions of canine excellence. Then in the 1960s the tip of the genetic iceberg emerged as concern grew about a joint disorder called hip dysplasia. A control programme involving the examination of hip xrays by a skilled scrutineer and the maintenance of a registry of animals “cleared” of the defect was established at the Ontario Veterinary College at Guelph, Ontario. Now after three decades of the OVC programme it has been pretty well established that “clear” animals with several generations of “clear” ancestry can nonetheless produce dysplastic progeny [Chidiac-Storimans 1995]! Hence the OVC control programme would seem to be of questionable effectiveness. As the generations of closed-studbook breeding have advanced, a panoply of other inherited problems has emerged in purebred dog breeds. There is no need to list them here; the list would be on its way to obsolescence in a month or so; veterinary research continues to define more inherited disorders regularly. Many breeders now run four-way screening programmes; some may screen for even more problems. Many breeders’ selection programmes for various kinds of canine excellence must now be at a standstill – all the selection is going into the effort to produce stock “clear” for eyes, hips, elbows, blood disorders, endocrine dysfunction, etc. Yet thirty years of xrays have not eliminated hip dysplasia – it is now widespread in breeds in which it was not a problem thirty years ago.
In December 1994, Time magazine published a scathing indictment of the American Kennel Club and of purebred dogs and their breeders, targeting in a cover story the problem of genetic ills, suggesting that the best use of pedigree papers was for housebreaking the puppies and recommending that the public satisfy its desire for canine companionship with mongrels. Since then, most of us have known we have an untenable situation on our hands. Our reputation as breeders of purebreds is now in tatters; we are caricatured in the media as greedy, uncaring producers of degenerate animals. The CKC’s main response to the situation was a Board policy statement that “reputable breeders will provide a detailed written guarantee of the present and future good health of the dog and will not hesitate to uphold their guarantees.” The policy statement, far from helping the situation, only saddled breeders officially with a heavy responsibility without enacting measures which might assist them in living up to it.
It is time for us as dog breeders to stand up for ourselves and for our dogs, to reject the imputation that we ourselves are individually to blame for the problem of genetic defects, and to demand swift remedial action by the Club and, if necessary, Agriculture Canada. The crux of the problem is the closed studbook and with it, the ideal of breed purity, the worship of type and the preeminence of the championship show as goal and arbiter of most breeding programmes. Armed with the concepts of population genetics, we can now examine the last: century of nineteenth-century dog breeding, ascertain what has gone wrong, and establish ways and means to correct the situation.
Earlier we stated that the recognition of a breed by a registry was a crucial event in its history, more crucial than it need be. That is because the usual practice has been to open the registry to foundation stock for a limited period, to inspect and register a small population of foundation animals, and then to close the registry to new genetic inflow forever after, with the sole exception of animals of the same breed imported from other registries and derived from the same or closely-related foundation stock. In recent decades there has usually been no unique Canadian foundation stock except in the case of indigenous breeds; CKC merely accepts registered stock from other jurisdictions. (Actually the relationship of CKC foundation stock to that of other registries has never been clearly defined, so far as I know. CKC accepts registration papers of other studbooks which it considers to be “reliable.” So long as the export pedigree shows three generations of registered, numbered ancestry; import stock seems to be eligible for CKC status without question. The criteria involved are clerical, not genetic.) Most of the breeds we are familiar with were founded from sixty to over one hundred years ago. In those days Canada’s population was much smaller than it is now; the canine population was correspondingly smaller, too. Thus the number of dogs accepted during the open-registry periods was rather limited.
The canine species possesses tremendous genetic diversity as a whole. Like most species, that diversity includes a genetic load, a wide variety of more or less deleterious alleles, probably quite a few of them held in a state of heterozygote superiority, so that although natural selection tends to eliminate homozygote recessives when they segregate, the bad alleles themselves maintain a strong presence due to the selective advantage of the superior heterozygote. What happens when a founder event occurs? Then it is possible that bad alleles, uncommon in the canine population as a whole, may achieve a much higher frequency of occurrence owing to their presence in a small founder population – especially since the foundation stock of a newly-recognised breed will already be considerable inbred from the breed development process. Inbreeding and selection together raise homozygosity levels dramatically through the wholesale elimination of alleles from the genome. Those alleles may be unwanted by the creators of a new breed; nevertheless their elimination raises the allele frequency of whatever remains.
An Example from One Breed
Thus the recognition of a breed creates a founder event when the registry is opened; a limited number of breed foundation animals are selected, often from a population which has already undergone considerable inbreeding and selection. Let us take for an example the Siberian Husky breed. Registered in 1939, the initial CKC population consisted of 47 animals, all belonging to or bred by one kennel! Of those 47, nine were foundation stock of the kennel whose dogs were registered. Two of those were males imported from Siberia – littermate brothers! The other seven were mostly related to one another. (Two were seven-eighths Siberian and one-eighth Malamute.) The other thirty-eight were all progeny and grand-progeny of the founders. Of the nine foundation animals, two were not bred from at all. Two were mated – once only – to each other: one only of their progeny contributed to further breeding. Of the two Siberia import males, one brother was always bred to the same bitch, producing a large number of progeny of identical pedigree; the other brother was usually bred to daughters of the first brother. Today Siberian Husky lines that trace directly back to the Canadian foundation stock owe 25% of their pedigree lines to the first brother, 15% to the second brother, and 27% to the first brother’s invariable mate! Two-thirds of the genetic heritage of these modern Siberian Huskies derives from only three foundation animals! This is not an exceptional situation, it is a fair example of the early breeding history of CKC breeds.
In the case of the Siberian Husky, then, (which happens to be my breed, with whose early history I am reasonably well familiar), The Canadian Kennel Club opened a registry in 1939, inspected one kennel’s dogs and admitted four dozen closely-related individuals to the registry, which was then closed permanently. No effort was made to ensure a broad foundation, nor a numerous one, nor a genetically diverse one.
Just how permanently the registry was closed I recently found out when I imported from Russia a dog bred to the Siberian Husky standard! The dog was born in the Ural Mountains well within the boundaries of Siberia from parents of Chukotkan village origins; he had three generations of known ancestry (without registration numbers since there is no official “Siberian Husky” registry in Russia). I was immediately told that the Club “did not know what to do” about my application to register the dog, that the protocols used to register breed foundation animals in 1939 were no longer valid, and that my dog “should not be used for breeding because it would probably be a long process,” in spite of the fact that the dog had a valid FCI Export Pedigree from the Czech Republic (through which he was exported). A year and a half later after repeated in camera discussions, the import was refused recognition by the Board and Registration Committee on grounds of inadequate information (no ancestral registration numbers). Repeated calls for Club inspection of the import and offers to submit the animal to DNA tests and progeny testing were ignored. The registry is closed – even to new Siberia imports!
For the past fifty-six years, then, all Siberian Huskies bred in Canada have stemmed from the 1939 registrations, or from American imports, which mostly stem from the same dogs CKC registered, plus perhaps three additional animals. The original foundation animals were poorly utilised and subsequent generations were so closely inbred that the two Siberia import males plus one bitch are even today still statistically equivalent to grandparents of every single Siberian now registered!
Thus the original founder event in my breed plus the closed studbook has resulted in a state of forced inbreeding for Siberian Huskies. There is no such thing as an outcross mating in Siberians in any genetically meaningful sense. A sire can be found, perhaps, who may have no ancestors in common with a bitch for the last 5 or 6 generations – if one knows all Siberian bloodlines well enough and doesn’t mind going a few thousand miles to find him – but he will not be an outcross, because all of his ancestors and all of the bitch’s ancestors are the same animals, once the pedigree is taken back far enough. It would be difficult to calculate inbreeding coefficients for fifteen to thirty generations of ancestry; software to handle calculations of that nature doesn’t seem to be generally available to breeders. (After all, a thirty-generation pedigree would contain over two billion names.)
Thirty generations of breeding all going back to ten dogs or fewer represents an impressive feat of sustained inbreeding! Predictably enough Siberian Huskies, which eighty-five years ago were probably the toughest, hardiest variety of dogs on earth, now suffer from the same gamut of genetic defects that afflicts other breeds. Few if any registered Siberians are now able to perform as sleddogs on anything approaching the level of the 1910 dogs imported from Siberia. Probably this is mostly due to the decline in heterozygosity and loss of vitality through inbreeding. What is worse, unmistakable signs of inbreeding depression are surfacing in the breed: rising numbers of Caesarean births, smaller litters, lower birth weights, delicate nestlings prone to infection, etc. Breeders of domestic livestock – cattle, poultry, sheep – manage to run registries and maintain breed type without imposing the concept of absolute breed purity. They inbreed to fix desirable traits, as do dog breeders. Livestock breeders, however, do not try to pretend that they can inbreed forever without ill effects. Thus when inbreeding depression or genetic defects threaten, they outcross – repeatedly, if necessary. They can do so because they do not have closed studbooks. I do not suggest that we slavishly copy the procedures and registry structures of livestock associations, because I think they, too, might benefit from some restructuring in the light of modern genetic knowledge. Nonetheless I would make the point that we in the canine fancy are in a minority when we cling to absolute ideals of breed purity and insist on rigidly closed studbooks.
As a dramatic contrast to the foregoing example of the CKC’s Siberian Husky breed foundation, let us examine for a moment the standards which Agriculture Canada now applies to new domestic animal breeds in this country, as set forth in a three-page leaflet entitled “Establishment of a New Breed of Animals in Canada.” Agriculture Canada now requires that breed foundation stock (that is to say, the first generation of registered animals of a new breed) be selected from the third filial generation (F3) or later of the “evolving breed” which precedes the actual, registered new breed. It lays down no parameters for the founder generation of the evolving breed, but it does state:
The standard used for the creation of a new breed is as follows:
- Minimum number of animals to constitute the foundation stock of the new breed (F3): 200 animals (unique genotypes).
- In order to reach the required 200 F3 animals and in order to provide a sufficiently wide genetic base, it is recommended that the minimum number of animals to be produced in each F level be:
- Fl : 60 animals
- F2: 100 animals
It also stipulates that “the F3 generation is the earliest generation to become eligible for inspection as foundation stock… In practice most evolving breeds will evolve over many generations before having developed a significant population of foundation animals.”
These modern standards are at least somewhat influenced by population genetics considerations, in an attempt to establish a basis for genetic health and stability for new animal breeds in Canada. Yet (in all probability very few of our existing CKC dog breeds, which are arguably of much greater economic importance than any new breed, would come anywhere near to the foundation stock numbers now enforced by Agriculture Canada. The sole exceptions would probably be breeds, like the Canadian Eskimo Dog, accepted for registration during the last decade or two. As for the Siberian Husky, its actual genetic founders (those whose genes contributed to future generations, leaving aside those which did not reproduce) numbered 6 only; the Fl generation which actually reproduced numbered 8 individuals; the F2 generation which actually reproduced numbered just 5 animals; no F3 animals were registered in the first year of CKC registrations — original founders. F1s and F2s were all registered together in the first year.
Thus it is obvious that the Siberian Husky, at least, could not begin to satisfy current Agriculture Canada standards for an appropriate number and variety of foundation stock to establish a new breed, when traced to its historic foundation. In all probability, few CKC breeds could do so. Yet the registry norms that are rigidly enforced by CKC, backed up by Agriculture Canada make the acceptance into the studbook of badly-needed new foundation stock a complete impossibility! Presumably Agriculture Canada has good and sufficient reasons justifying its standard for new breeds — that being the case, then it is a curiously irrational situation that older, existing registered breeds not only are exempt from any such standard, but are actually prohibited from enlarging their founder group by the acceptance of unrelated primitive stock.
The Holistic Breed
Now I would like to evoke a vision of the future — but not the distant future. I want to describe how dog breeds might be in the twenty-first century. Instead of all breeds being subjected to arbitrary structures not equally well suited to them all, each breed would get whatever special measures its breeders thought necessary. Instead of a fragmented canine fancy with ghettos of show, fanciers, obedience buffs, and working-dog specialists, dog breeds would have the benefit of a holistic outlook, integrating the various aspects of canine: activity and producing well-rounded, versatile, mentally stable animals. Let me stress that the suggestions which follow will be fully practical and down-to-earth. They involve no technology we don’t already possess. They require no knowledge that isn’t already generally available. All that is needed is a proactive attitude and the will to make necessary changes in an obsolescent structure. This vision could become a reality within ten years time.
At the beginning of this brief I stated that the three distinct axes along which breeds are distinguished – ancestry, purpose, and typology — had to relate fully and cooperatively, or the fullness of breed identity would be missing or marred. Let me now describe how such a relationship might he achieved.
To begin with, we absolutely must open CKC stud books, in every breed, to new genetic inflow. There can be no long-term genetic health in small populations such as our registered breeds without the periodic infusion of new genetic material. The one big “sacrifice” we shall have to make, if it is really a sacrifice, is to abandon racist attitudes and the concept of rigorous breed purity. We must recognize that first of all, a dog is a dog, species Canis familiaris, and that is his true identity. He is a dog first, before he is a Siberian Husky or a Foxhound or a Doberman; breed identity is subordinate to species identity. We must stop treating breeds as if they were species, abandon the rigidity and narrow typological thinking which has heretofore characterized the canine fancy. We must recognize that dogs are unique individuals and that there is no positive value in trying to create groups of dogs which are all clones or photocopies of a type specimen represented by a breed standard. This should not be too hard, since breeders and judges have never been able to arrive at agreed and consistent interpretations of breed standards anyway. Why, then, should we pretend that a standard, which as it now exists evokes a different imagistic interpretation in the mind of each individual breeder and judge, describes a single ideal type?
Canine breeds can and should be differentiated, bred and maintained on a dynamically balanced, heterozygous population basis without restriction to a closed, historic founder group. The closed studbook and the breed purity concept are, from a genetic point of view, simply unnecessary. Indeed, as we have seen, from the standpoint of maintaining a genetically healthy limited population, they are thoroughly counterproductive. Where is the logic in submitting each and every CKC breed to a registry system which guarantees ongoing, progressive genetic degeneration, loss of species vigor and hardiness, and saddles every breeder with the unwanted, unhappy responsibility of producing more and more unhealthy, flawed stock as time goes by? The notion that genetic disease can be controlled, much less eliminated, by screening programs and selection has not been borne out by general experience. Those who promote such a notion are engaging in a cruel, self serving deception. It may be that a breeder can sometimes improve his odds against producing defective stock in a given mating by screening the parents, but experience has proved that screening will not solve our genetic problems in any wider sense. Despite generation after generation of “clear” stock, bloodlines can still produce more and more affected animals. That is because our problems are inherent in the closed studbook/incest breeding system. In order to restore genetic health we shall have to adopt a different system.
It will be asked, “Just how will the opening of our studbooks to outcross stock bring about the elimination of genetic defects?” The answer is that it will not eliminate genetic defects. That need not be the end in view. If we could somehow eliminate all the various genes now known to produce harmful anomalies, plus all of those yet to be discovered, we would almost certainly find that the remaining genome was non-viable, that healthy reproduction and growth to maturity could not reliably take place. Genetic defects are not “eliminated” in nature. Instead random mating and behaviour patterns that discourage inbreeding take care of the problem by ensuring high levels of heterozygosity and the consequent rarity of defective homozygotes. If we take steps to set up similar patterns in purebred dogs, we shall be able to reduce the level of expression of defective genes greatly, which is all that is required. The end in view is healthy stock, not “racial purity.” Purged and purified bloodlines would be weak for other reasons, as has been explained. As the mapping of the canine genome progresses and RFLP allozyme or microsatellite “markers” for common genetic defects are found, we shall probably then be able to use DNA studies to recommend matings that will avoid the production of defective homozygous progeny — provided that we have made enough genetic diversity available through outcrossing to give us the genetically distinct lines from which to choose! As things stand now, most breeds are so homozygous that it could prove extremely difficult to find matings which would avoid one genetic defect without reinforcing another!
New Structures for
the Dog Fancy
Very well, then, if we eliminate the closed studbook, how shall we decide what stock to admit for registration? One must begin, of course, with the existing body of registered stock. Thereafter, one way of proceeding might be to strengthen and empower the breed clubs. They should be granted responsibility and autonomy: responsibility for the welfare of their breeds, and autonomy to make the judgments and decisions necessary to fulfill that responsibility. It should also be ensured that the breed clubs are fully representative of all breeders, by making breed club membership a requirement for anyone wishing to register stock he has bred or imported.
The first task of the breed clubs would then be for each of them to determine what sources of genetic inflow might best be employed in their breed. Breeders alone can command the collective expertise to make that decision and it ought to be theirs alone, but the designation of outcross sources should be obligatory, not optional. The Siberian Husky Club of Canada, for example, would have to decide where outcross animals might best be obtained for restoring heterozygosity to that breed; they might decide, for example, that dogs imported from Russia and perhaps even an occasional outstanding individual carefully selected from the present “Alaskan husky” gene pool of racing sled dogs (which was derived largely from 1910-era Siberia imports that remained in Alaska) are two logical sources. Breeds which do not have their origins in autochthonous populations would have to seek outcrosses in similar related breeds, as Spaniels (English Springer) and Spaniels (Welsh Springer), or Retrievers (Labrador) and Retrievers (Flat-Coated). They would then have to set up inspection and test-breeding procedures for admitting outcross animals. Once the outcross sources had been designated, selection of candidate animals would in most cases be best left to individual breeders, who might then apply to the breed club for preliminary inspection of their outcross — which inspection ought not to be excessively rigorous. General soundness, reasonable temperament, proven working ability, approximate size and physique, and acceptable overall type should be adequate criteria, none of the foregoing to be rigidly interpreted. The outcross should then be registered provisionally by CKC, subject to breed club inspection of two generations of its progeny. The registry should remain permanently open to new outcross animals. It might prove desirable to set limits to the number of outcross dogs registered in any given year proportionate to the overall breed population, in order that small populations not be swamped by excessive outcrossing. Some regulation of the process would obviously be necessary, but the best regulation would probably be breed club oversight and guidance of the process, backed up by CKC supervision.
Advantage should be taken of DNA analysis techniques by using them to monitor heterozygosity and relative kinship in major breeding lines. (It would also be a good idea for the Club to offer DNA profile parentage certification.) This technology already exists and is in use: it is rapidly becoming much more affordable. Limits should definitely be set on inbreeding, preferably by the breed clubs, but CKC should decide maximum allowable limits of inbreeding as a default setting. Only by the outright prohibition of excessive degrees of inbreeding will it be possible to make the transition to a balanced-heterozygous state for purebreds; otherwise old ways will continue through inertia and persistent typological thinking. Assortative mating can and should become the norm for the preservation of type, mating individuals which are phenotypically similar but unrelated or at least not closely related. The Club would have to monitor registrations, possibly performing occasional DNA spot-checks, to ensure that inbreeding does not take place; otherwise many would continue to breed from whatever dogs were in their own backyard rather than seeking breed club advice to find suitable individuals from unrelated lineage.
A Healthy
Balance, for Breed Identity
The responsibilities of the breed club should not end with the designation of outcross sources and the inspection of outcross candidates. If the fullness of breed identity is to be achieved overall in each population, then the breed clubs should take on responsibility for balancing the various facets of that breed identity. Realistic, meaningful and workable systems should be introduced for monitoring temperament, for proving working ability and trainability, and for evaluating, type and appearance. Championship shows would then become breed-club events, since the methods of evaluation and the various events required to test such qualities as temperament, vigor and endurance, working ability, and trainability would be breed-specific and under the breed clubs oversight. That is not to say that a number of breed clubs might not band together to stage events for several breeds simultaneously at the same venue, but the all breed show with all-rounder judges, under CKC rules for CKC Championship points, would eventually be history. To ensure wholehearted support and participation by breeders, it would probably be necessary for CKC to evolve some means of making clear on the papers of every dog the extent to which that animal had been submitted to the testing and evaluation procedures of the breed club and with what result. Breed club input of information to the Clubs database could be done by e-mail on the day of the event. Strong incentives for participation should be arranged and breed clubs should be so structured that they could not be autocratically ruled by individuals or cliques.
Registration certificates produced by CKC would carry much more detailed information under the new system than they now do. The computer power is now available to make this quite feasible. A certificate of registration should once again carry a pedigree of at least four generations. A two-tier certificate system would be necessary, as no dog would be eligible for breeding registered progeny until it had been inspected and evaluated by the breed club. Rating and measurement protocols are already being worked out by proponents of the Advanced Registry proposal. Broodstock certificates should carry a summary of the breed clubs rating and evaluation of the animal, together with evidence of proof tests for temperament, working ability and trainability. All certificates should identify outcross lines and bear a quantitative estimate of the relative heterozygosity of the animal identified by the certificate.
Breed standards would require revision under the new system. The concept of disqualifications should probably be dropped in favour of a detailed rating system in which all breeding stock would participate. In the case of quantitative characteristics such as height and weight, a simple Bell curve statistical description of the desired mean and range ought to be sufficient, without disallowing occasional extreme examples. Working abilities ought to be clearly defined in the breed standard and a basic performance standard given where possible. Clearer and more detailed descriptions of desired temperament and of qualities bearing on trainability ought to be part of the new standards. Prescriptive minutia should be minimal; it should be sufficient merely to describe the general distinguishing features of a breed, without an excess of cosmetic and conformation restrictions, except where indispensable breed points are involved. Type stringencies should be relaxed considerably, allowing most breeds to become moderately heterotypic; if qualities of working ability, hunting instincts and similar traits achieve greater emphasis, there will be correspondingly less need for extreme type requirements to distinguish breeds. Standards should be holistic descriptions of the breeds they identify, brief statements of essential breed qualities, rather than typological blueprints. It is imperative to subordinate typological thinking to considerations of utility, genetic health and hardiness. First a dog should be healthy, balanced, of sound mind in a sound body, able to fulfill his breed purpose; after that can come points of beauty and type but never again in the bizarrely exaggerated fashion that now prevails in the breed rings of championship shows.
It might eventually be found desirable to quietly merge scarce and consistently unpopular breeds, as well as closely similar breeds, with populations nearest to them in general characteristics, possibly initially designating them as breed varieties. Reasonable numbers are necessary for the maintenance of a healthy population. The number of breeds recognized has continued to grow, yet the total number of dog owners in the country may not have grown proportionally. A rare breed is not the same thing as an endangered species; breeds can come and go without damage to the canine species as a whole. Breeds known to be of low viability due to their dependence for breed identity on anomalies such as achondroplasia, may have to be dropped from the registry unless evidence is advanced that they can be upgraded to certain minimum standards of health and structural soundness.
Can it Really Work?
I can hear someone objecting, after having thought about the idea of a breeding and registry system in which outcross breeding was actually encouraged, “Surely this system will produce some dogs which are not even recognizable representatives of their breeds! What happens then?” Typological thinking dies hard. I used to worry lest my Siberian breeding program should one day produce a dog or dogs whose ears were not fully erect. It never happened. Instead something much worse happened when I found that I was producing some dogs who ran a high risk of being unable to lead a healthy, normal canine existence, through endocrine malfunctions, immune system weakness, and the risk of blindness. To think I had worried about the possibility of a tipped ear, something which would not handicap or bother the dog in the least! Let me say the following, then, to those who worry that a balanced-heterozygous breed will engender “untypical” examples. It is far better that our breeding occasionally engender a dog deficient in breed type, than that we should consistently produce large numbers of dogs guaranteed to lead lives of suffering, creating anxiety, large veterinary bills, frustration and unhappiness for their owners. That is what we are doing now. Over sixty percent of Golden Retrievers, for example, will suffer from hip dysplasia, osteoarthritis or osteochondritis in their lifetimes. Is that to be preferred to the possibility of producing an occasional robust “mutt” lacking in breed type but who will nonetheless still make someone an excellent happy, healthy companion? I am sure that it would take awhile for all of us to learn how to breed in this new and different way; I suppose we might produce occasional oddities in the process. Yet I am absolutely convinced that the good results we would quickly achieve would more than make up for the embarrassment of our failures. At the very least we should all have clean consciences once again, knowing that we were making our best efforts, using up-to-date genetic knowledge, to produce sane, healthy, robust canine companions. Let us not forget that as DNA mapping procedures advance (there are at least two canine genome mapping projects now under way) our tools are going to improve and our ability to predict what our breedings may produce will be greatly enhanced.
As things now stand, the dog fancy is in a position which is frankly untenable. The CKC Board of Directors has unilaterally committed “reputable breeders” to the proposition of guaranteeing the “future genetic good health” of the dogs they sell. Yet those same breeders have no means of protecting themselves from the looming specter of financial ruin should they be held to such a guarantee, otherwise to the loss of public credibility. Other than the continued elaboration of screening programs and the Advanced Registry proposal, both of which are somewhat like applying an adhesive bandage to a severed artery, nothing is being done about making guarantees of genetic health a workable proposition. At present, purebred breeds — all breeds — show levels of genetic defects totally inconsistent with the practical maintenance of the Boards policy. Many honest, caring breeders are racked by torments of guilt and self-reproach brought on by the suffering of defective dogs, yet it is really no fault of the breeders themselves! The fault, as has been demonstrated in this brief, lies with the closed studbook and the inbreeding system. If the consensus of the Club is truly that purchasers of purebred dogs have a right to expect genetically healthy animals, then the Board has no choice other than to do everything in its power to change the existing system so that healthy animals may once again be reliably produced! That will never happen just through Advanced Registries, higher Championship point requirements, more screening programs, and Board policy pronouncements. The Club must take to heart the lessons of population genetics. It must open its studbooks to outcross stock on a permanent basis. It must take measures against the obsessive pursuit of breed type and the worship of breed purity, measures which will increase the health, utility, trainability and sanity of purebred dogs, measures which will balance the elements of breed identity. There are no credible “soft options” left.
One unfortunate reality which must be faced, however, in order to bring about any major changes involving the CKC will be the conservatism and resistance to change of the Board and of the “old hands” — the ruling oligarchy of the Club. The CEO and the Board will almost certainly aggressively defend the status quo no matter how urgent the need for change. At present, for example, they turn down requests for the registration of new foundation animals with statements such as this one: “The CKC takes pride in registering dogs based on accurate and complete information and we will continue to strive for these high standards.” Yet when that statement was written, the Club was still registering Canadian-bred litters whose parentage information was supported only by a signed registration application form filled out by the owners of the dam and sire. Under that system of information gathering it is regularly necessary for the Board to cancel litter registrations when it becomes evident that the parents of some litters are not both of the same breed. No one knows how many litters go unchallenged which, although purebred with both parents of the same breed, have their parentage misrepresented because the actual sire of the litter is not the dog entered on the application form. In the absence of DNA testing, how can the substitution of sires be detected?
Meanwhile the United Kennel Club, a “dissident registry” in Kalamazoo, MI, USA, which now registers about a quarter of a million dogs annually, has already instituted a process for the verification of parentage by DNA profiling! This is the first time that DNA profiling has been made routinely available to dog breeders, and UKC is the first canine registry in the world to offer such assurance of verified parentage. Innovations such as this make the Clubs defensive statements about its high standards sound rather hollow.
Anyway, those of us who seek reform will have to contend with a Club establishment which will attempt to make a virtue of the very things which most threaten the genetic health of CKC dog breeds: the closed studbook, the breed purity concept, the endless inbreeding, the constant refinement of type, the preeminence of the Championship show, Those who dare to challenge the existing system will have to put up with being made to look foolish or even villainous by the solemn pronouncements of the old guard. We should all realize that the Club establishment is unlikely to initiate serious action for change in the absence of grassroots pressure from the general membership. It is up to us to initiate serious dialogue along the lines outlined in this brief, to research ways and means to promote a different, healthier method of purebred dog breeding, and to raise the consciousness both of novices and of old hands regarding the genetic dilemma which now faces us.
Deep structural change cannot occur without widespread debate among fanciers, because new and different concepts sound threatening when they are first described. Once the reasoning behind them has been adequately discussed, the threat often disappears. Someone may ask, for example, “What about these open-ended Breed Standards? A Bell-curve statistical description of a breeds height standard may be an adequate formula, but what if the mean is set at 22.5 inches and you don’t disqualify the 25-inch dogs. Then maybe in a few years the mean may drift upward to around 24 inches, with hardly a single dog under 22 inches. What then?” My answer would be that the whole point of the balanced-heterozygous system is its healthy flexibility. A stubborn insistence on narrow tolerances in matters such as height at withers usually involves the sacrifice of other worthwhile qualities anyway, as too many otherwise good animals must be discarded only because they are a shade over standard. In the balanced system described, nothing at all need be lost. If the height mean of a 22.5-inch breed should drift upward to 24 inches, it would be because most of the breeders wanted a taller dog! Since the breed club would be advising breeders, measuring and rating dogs, maybe even suggesting matings, this sort of gradual change would occur only with the knowledge and acquiescence of the breed club, representing all active breeders. Under a heterozygous plan with mainly assortative matings, nothing whatever is lost in such a gradual change. Should the height drift upwards and, later on, the breeders decide upon a return to the original mean, a simple shift in the emphasis of assortative mating will accomplish such a return easily, smoothly, with no genetic loss and no disturbance of other traits.
The whole idea of a dynamically balanced heterozygous breeding system is the retention of as much healthy genetic diversity as possible. Such diversity makes it easy for a breed to develop and progress in whatever direction its breeders wish. It also ensures that genetic problems are kept to a minimum no matter what changes of standard may occur. In the statically balanced homozygous system now in force, the more homozygosity increases with time and selective breeding, the harder it becomes for major change to occur naturally and easily, and the more pronounced genetic problems become. Once an allele has been “fixed” in homozygosity, no amount of selection can change that trait; only radical outcrossing can restore the lost alleles and such outcrosses will always upset the static balance completely, necessitating years of remedial inbreeding and selection, probably creating new genetic problems. I am convinced that a system based on a dynamic equilibrium of healthy dominant genes must inevitably be better than one which throws away most of the healthy genetic diversity in order to achieve static stability for homozygous recessive traits.
It is worth noting that the new system, if carried out at all conscientiously, would mean more work per dog for everyone. Breeders would necessarily invest more time and effort in their breeding stock in order for it to pass breed club requirements. This is by no means a negative factor. One ongoing problem in our society is that of large numbers of unwanted pets. Another related problem for the purebred fancy is substandard dogs produced by the non-serious “backyard” breeder and the puppy-mill profiteer. The suggested reform measures would discourage exploitative factions and reduce considerably the overall number of purebred dogs, while raising greatly the overall quality levels and ensuring that practically all purebred dogs were valuable, cherished, and wanted by their breeders and owners. The new system would greatly increase the inherent value of purebred canine stock. Purebred would then mean much more than just a paper certificate!