A COMMENTARY ON THE COA CROSSBREEDING PROGRAMME
From Chinook Owner J. Jeffrey Bragg

 

Commmentary written September 2010

Arthur Walden with "Chinook"

Recently a Chinook genetic study by Mars Veterinary corporation was made available on the Chinook Owners Association website. The results of that study have serious implications for the COA Chinook Crossbreeding programme, not so much due to the revelation of new information as to the underlining of basic genetic principles that may not have been given adequate consideration in the design of the programme. Probably the most significant statement in the report is: "If it was a goal of the cross-breeding program to increase the amount of traits from other breeds or crosses to the Chinook, we would suggest that these were introduced to the purebred population at an earlier generation before the majority of added genetic material was lost." I might add that the same observation holds true if the goal were to increase genetic diversity in the Chinook.

Defining Terms

I shall use some very basic genetic terms in this message, so I'll define them here just to make sure the message can be clearly understood by all.

Allele - a specific version of a variable gene. A single gene or section of DNA that controls an inherited trait may have several different alleles or versions. Only one allele, however, can be transmitted to an offspring by each parent, with respect to that particular gene.

Homozygosity - the presence in an individual's genetic makeup of identical alleles with respect to a given gene, meaning that the same allele was transmitted by both the sire and the dam. Similarly, the term is used in a more general way when discussing the overall proportion of an individual's genetic heritage that is thus identical, derived from both parents.

Heterozygosity - the presence of non-identical alleles with respect to a given gene, meaning that the sire and the dam have transmitted different versions of that gene to a particular progeny. Also used more generally when discussing the degree to which an individual's genetic heritage is diverse, with different alleles inherited from each parent.

Inbreeding depression - loss of vigour, hardiness, reproductive viability and immune system strength in inbred bloodlines and individuals, through the expression of lethal, sublethal and suboptimal genes, as well as by the loss of overdominance benefits.

Overdominance (heterozygote superiority) - fitness advantage possessed by heterozygous individuals, displayed in greater vigour, hardiness, reproductive viability, immune system strength, etc.

Inbreeding coefficient - A percentile or decimal figure derived from counting the repetitions of individual known ancestors in an animal's pedigree and summing such inbreeding up by the application of a formula. This can't be done by hand for long, complex pedigrees, so COI is always calculated by computer software. The resulting figure may be taken as indicative of the probable proportion or percentage of genes that will be homozygous by descent for a given mating, or equally well as the probability that any given variable gene will be homozygous by descent in that mating.

Inbreeding in Chinooks

I would question whether sufficient attention has ever been paid to the subject of inbreeding and its consequences in the Chinook breed. Part of this may have to do with the fact that Chinook pedigrees are not easy to pin down. There are too many gaps, too many "open pedigrees" and disputed points of ancestry. In some cases these gaps simply cannot be filled because the ancestry is not known; in others, the knowledge exists but is disputed by some and therefore ignored by most breeders. The situation is further complicated by the history of the breed. The loss of pedigree continuity in the Harold Smead period means that the ancestry of the 1981 rescued founders of the UKC registered breed cannot be joined up with the known pedigrees of the Wonalancet-Hubbard period. This in itself has serious consequences that have been largely ignored.

In our own Atholl Chinooks database as installed in Breeders Assistant (a pedigree record keeping application that also calculates COI), we have filled in all of the gaps for which reasonable data exists; I refer to Northwind Nome, Northwind Tia and a few others. With the known data plugged in, that database generally indicates 10-generation Chinook COIs on the order of 20% to 30% or more. (For those who don't know, this is a pretty high level of inbreeding -- the COI of a brother/sister mating or a father/daughter mating is 25%.) Trying to get a handle on the present overall degree of Chinook inbreeding, we input to our database a recent five rounds of "New Arrivals" from the COA Quarterly. The lowest 10-generation litter COI was 30.43%; the highest was 40.04%; the mean was 35.24% and the average over 19 litters was 35.48%.

The higher the levels of sustained inbreeding and the longer such inbreeding is carried on, the greater will be the homozygosity in the subject population -- in this case, the Chinook breed. For many decades of the 20th century it was believed that homozygosity was a great virtue. Showdog breeders believed in the breed standard as a pattern for perfection. Although people's interpretations of standard varied, show judging provided enough consistency to result in the development of a "cookie-cutter" similarity among individual showdogs; this was achieved through high levels of inbreeding and selection. It took far too long for the truths of population genetics to reach the dog world. Even today, long after zookeepers, wildlife biologists and other animal husbandry professionals have accepted that genetic diversity must be a primary objective in population management, the majority of the kennel-club showdog world has yet to receive, understand and accept that message.

Sustained inbreeding to achieve close conformity to a single desired phenotype will result in high levels of homozygosity, together with a corresponding lack of genetic diversity, resulting in a breed genome much like that of the cheetah, totally lacking in genetic variability, or (as geneticists describe such a genome) "depauperate." This lack of diversity will be observed not just in terms of superficial cosmetic traits, but also in critical fitness traits such as the Major Histocompatibility Complex of genes responsible for immune system strength. High levels of homozygosity will also affect the genetic load, which means that recessive genetic diseases will find increased expression among individual animals. Chinook breeders tell themselves that "everything is all right because we have been assured by the Mars study that our breed is 'in the middle' of purebred breeds in terms of diversity." That assurance is not worth much, because scores of purebred dog breeds are also in the same situation as the Chinook, depauperate, having suffered from the same breeding practices for a century more or less; the difference being that most such breeds have far larger breeding populations than the Chinook, hence a better shot at survival. Chinook breeders seem to take comfort in the fact that 20% and 30% Coefficients of Inbreeding are not uncommon in other dog breeds. But is it really true that the Chinook is no more inbred than that?

Just How Inbred Are They REALLY?

Arthur Walden's "Chinook"

Unfortunately I do not for one moment believe that these calculations fairly represent or fully reveal the true extent of cumulative inbreeding in Chinooks. We know that when Arthur Walden returned from Antarctica, had his falling-out with the Seeleys, and started Chinook breeding over again with Julia Lombard, the foundation stock for Wonalancet-Hubbard kennels consisted of just three animals: Jock, Zembla and Hootchinoo. Jock and Zembla were brother and sister, sired by Walden's leader "Chinook" out of a Belgian Shepherd female; Hootchinoo was sired by Chinook out of his own daughter from a German Shepherd bitch. Since Julia Lombard's avowed objective was to create a purebred Chinook, it is reasonable to assume that during the ten years of Wonalancet-Hubbard breeding, no further crosses were introduced. Pedigrees of some Wonalancet-Hubbard litters show Jock bred to progeny of Hootchinoo and Zembla with 56.25% percentage blood of Walden's lead dog. Perry Greene Kennel bought the Lombard stock and continued to breed on the same foundation, having contracted with Julia Lombard to breed Chinooks only as purebred and to make no outcrosses without her express permission. For how many generations and on what plan the Perry Greene inbreeding continued are facts that seem not to be generally known or adequately appreciated by today's Chinook breeders.

My wife Susan has been reviewing her own research materials and memorabilia, including a number of handwritten and typed vertical family-tree pedigrees of Perry Greene dogs, some of them on the kennel letterhead. She has now entered these into Breeder's Assistant and come up with some interesting results. By the middle to late 1940s, several dogs in these pedigrees were already at COI levels in excess of 50%! Unfortunately we do not have enough data to connect this body of pedigree information with the Sukeforth rescue animals, making it impossible to arrive at an accurate COI for the 1981 breed founders.

A parallel to Chinook breeding, however, may be seen in the case of the dogs of Jacques Suzanne, another gentleman dog breeder and adventurer on the Walden plan, who bred his own special strain of Siberian Huskies in Lake Placid, NY for many years and then turned his breeding over to Dave Irwin, who carried on breeding the same pure bloodline -- all of them AKC registered and all descended from the mating of two dogs owned by Suzanne in the late 1920s, Polaire and Darka. I have been able to track the Suzanne/Irwin breeding down to the mid-1960s. By that time the COIs of the Suzanne bloodline had reached figures on the order of from 66% to 72%. I think we have every reason to believe that the 1981 rescue dogs probably had very similar COIs. Therefore, treating the earliest known ancestors of PG Riki, PG Barrow et al. as unrelated founders is totally unrealistic. Late Perry Greene breeding showed abundant evidence of further inbreeding, with multiple repetitions of PG Trondek, PG Attu I, PG Kim, PG Atka et al. If we could connect up the gaps of the Harold Smead era I believe not a single purebred Chinook of today would have a whole-pedigree COI of less than 50% and probably the figure would be on the order of 70%. Small wonder Harry Gray called these dogs "the walking dead."

Questions We Should Ask

Anyone contemplating the breeding of a Chinook litter should ask herself some searching questions:

- As a Chinook breeder, do I actually WANT to own and breed dogs with a 70% Coefficient of Inbreeding?

- Do I actually approve of this kind of breeding?

- Does a severely inbred animal represent my ideal Chinook?

- Do I feel that extremes of this sort are necessary and justifiable in the service of my (or anyone else's) notion of breed type and breed purity?

- Could it be possible that all the Chinook health concerns that are discussed every day on the Chinook email lists could have been brought on by these levels of inbreeding?

- Might it not even be probable that this is the case?

- Do we really have to subject our dogs to this in order to have a breed, or could we reduce the inbreeding to less than 10% and still have Chinooks -- HEALTHY ones?

Is Inbreeding Easily Avoided? (A Demonstration)

Atholl Charles Edward Stewart, one year old

I would like to offer a small demonstration of how difficult it is in the present circumstances to avoid inexorably increasing levels of inbreeding. To do so, I shall make some very liberal assumptions. (If these assumptions are unrealistically liberal, the fact will only strengthen my case.) Let's assume that the COA Crossbreeding Programme Committee accepts my Seppala female Darka of Seppala as a Dog Zero, with Tullibardine Duke Ellington as her mate. This mating actually occurred and we kept four progeny. Let's assume that Susan and I also breed our accepted Chinook Cross (81% G4) male Tullibardine X Holunder B-15 to purebred Tullibardine Queen Moussette. Now let's assume that Atholl Charles Edward Stewart, the male from the Duke/Darka litter, is put to purebred bitch Atholl Odetta. (I know she's too inbred, so I would want to mate her with as unrelated a boy as our kennel affords.) Let's assume that a few years downline we mate a boy from the Charlie/Odetta litter to a girl from the Lander/Queenie litter. And finally, let's assume that, hoping to conserve the genetics represented by ancestor Tullibardine Power, I finally mate his son Tullibardine Fram to both a Charlie/Odetta female and to another female from the Lander/Queenie x Charlie/Odetta litter. Here are the 10-generation COIs for all the dogs and litters involved:

Atholl Odetta (purebred Chinook) 48.02%
Tullibardine Queen Moussette "Queenie" (purebred) 40.75%
Tullibardine Fram (purebred) 40.75%
Tullibardine Duke Ellington (purebred) 35.12%
Tullibardine X Holunder B-15 "Lander" (grade Chinook from Maverick outcross line) 21.08%
Darka of Seppala (Seppala Siberian Sleddog, granddaughter of Siberia import) 16.83%
Atholl Charles Edward Stewart "Charlie" (F1 outcross) 0%
Lander/Queenie progeny 25.70%
Charlie/Odetta progeny (F2 backcross) 21.03%
Lander/Queenie x Charlie/Odetta litter (F3 generation with added Maverick outline) 27.73%
Fram x Charlie/Odetta litter (F3 backcross) 36.18%
Fram x Lander/QueeniexCharlie/Odetta (F4 backcross with additional cross line) 37.84%

Note that the progeny of Duke and Darka have a 0% COI, as there are no known ancestors common to both sire and dam, hence no known inbreeding! The most inbred animal in the group above is little Odetta, offspring of a grandson/granddam mating out of old Tullibardine Mustang (by Northdown Ootah ex Perry Greene Natanis), who herself had only a 28.44% COI; but Odetta's known COI is quite high at 48.02%. So, despite the fact that Charlie is a 0% COI F1 outcross progeny -- not inbred at all as far as can be told from the known pedigrees of both parents -- as soon as we mate him with a Chinook purebred the Coefficient of Inbreeding returns immediately to over 20%. Lander (by Mountain Laurel Bowerbank BB ex Singing Woods Crossing Mollyockett), who has the Siberian Husky outcross as his maternal great-grandsire, has a relatively moderate 21% COI, but his progeny out of purebred Queenie would have 25.70% COI. Now you might think that crossing these two outcrossed litters, Charlie/Odetta and Lander/Queenie, might give us some relief from the inbreeding; not so -- the resulting COI is another step higher at 27.73%. And if we breed purebred Fram (Queenie's brother) to a bitch from the Charlie/Odetta litter for an F3 backcross generation from the Duke/Darka outcross, we are almost back where we started, with a 36.18% COI, only three percentage points short of Fram's own purebred COI. The result of the mating of Fram with an F3 bitch progeny of the two outcross litters is yet more discouraging; even adding the Maverick outcross line, the COI is still 37.84%.

The lesson from this series of trial matings is clear: even though we WANT to reduce the COI by introducing a new Chinook Cross lineage, we cannot accomplish the goal! With only the second generation we are immediately back to the 20% level, and F3 and F4 backcross generations will be over 30% COI again, even if we were to combine our two outcross lines. The three-generation backcross requirement totally defeats our goal.

Now someone might ask the question, what would happen should the Maverick crossbred Chinook Holunder X B-15 be mated to a female of the Duke/Darka litter? (Under the current rules to do so would probably penalise us by adding one more generation to the upgrade process, so far as we have been able to determine.) But IF we should try it anyway? The COI of a litter sired by Lander out of Charlie's sister, Atholl Flora Macdonald, would be only 13.30%. And at long last we would be headed in the right direction to reduce those sky-high levels of inbreeding. It is perhaps a possibility that some of the progeny might turn out somewhat husky-ish in appearance. Would that be a disaster? Would it be anything that an intelligent breeder could not manage through selection? I don't see why it should.

Breed Purity or Genetic Health? Which is it to be?

What are our values? Do we value genetic health over "breed purity"? Just what does the phrase "breed purity" mean anyway in the context of a breed like the Chinook, which was a mongrel right from the get-go? What magic turns a mongrel into a purebred? Relentless inbreeding? If so, is it really worth it, if the resultant animals are riddled with health issues?

Stripped of all the discussion of individual dogs, individual breeders, and individual genetic problems, the real issues are rather simple and crystal-clear. Decades of unrelieved inbreeding strip any breed of its genetic diversity and, combined with founder effect, cause multiple "genetic defects" to be expressed in the phenotype of the resulting animals. Most of these components of the "genetic load" (geneticists prefer this term to "defects") are recessive. Most of them have very low gene frequencies in normal, healthy, outbred canine populations; they simply are not a major problem. But when a handful of founders happens to have several such genes, the gene frequencies in the resulting population can be very much higher; repeated inbreeding/selection cycles then make the situation much worse through random drift. Sustained inbreeding results in inbreeding depression. It also removes the hardiness that stems from overdominance or heterozygote superiority.

Nature knows and cares nothing about breed purity. That's an artificial human concept. Come to that, she knows nothing of "breeds." All animals of the same species can interbreed with fertile results; breed barriers are entirely artificial and exist only in the minds of the breeders, they have no basis in reality. Purebred dog folk talk about breed purity as though it were some kind of exalted natural virtue. It is nothing of the kind. It is something comparable to various historic discredited notions of racial superiority and perfection, such as those of the eugenics movement of the late 19th century.

It is fully feasible to breed and select dogs within a defined canine population without drastic inbreeding and still come out with a group of dogs that have characteristics and traits desired by the breeders. What is not possible, however, is to close that population to continued fresh gene inflow and still avoid inbreeding and inbreeding depression. Genetic health will inevitably be negatively affected; it takes awhile for the effects to become egregiously obvious, and by that time it is TOO LATE to fix the situation with timid half-measures like those of the present COA CB programme.

Solving the Inbreeding Problem

J. Jeffrey Bragg and "Charlie"

The solution is fairly obvious: introduce more Dogs Zero, and release the resulting lines to interbreed with the overall Chinook population much earlier in the game, at the F2 level. The Mars study makes very obvious that survival of new genetic material beyond the F2 level is far from guaranteed, and that very little indeed is likely to survive to the F4 level when strict selection to the breed standard is applied at each generation. As concentrated as the Chinook genetic heritage is (as demonstrated by the strong clustering shown in the Mars report), I would expect that to maintain an overall level of 50% contribution of the old Chinook purebred lineage would be all it would take to ensure the continuance of unique Chinook appearance and personality traits. If most of the Dogs Zero were to come from established working sleddog lineage, such a programme might have the added benefit of considerably enhancing working ability overall.

What is quite obvious, at the minimum, is that the existing Chinook CB programme needs to be radically revamped. In its zeal to ensure healthy stock and to defend the "purebred" Chinook label, it sets the bar so high that it fails to accomplish the desired result of enhancing Chinook genetic diversity. The guidelines that somehow morphed into requirements are so stringent as to exclude most if not all candidate Dogs Zero (or more realistically, Chinook cross F1 litters, since in practice nearly all such matings are accidental rather than planned, perhaps not surprisingly in the light of the discouragingly high requirements). For the genetic health and welfare of the breed the existing Chinook Cross programme really should be replaced with a more open and minimalist programme to encourage the development of more numerous Chinook cross lines, and allow such lines to be released for unrestricted breeding at the F2 generation when they can still make a meaningful contribution to Chinook genetic diversity.

It astonishes me that no one appears to have stood up and made these points to COA and its breeders long before now. There is little that is abstruse, difficult, or questionable here. The genetic principles involved are accepted and quite well known. Nonetheless I imagine that some, at least, might question my own qualifications to say these things so forthrightly and authoritatively. Anyone who does not believe me is welcome to ask Dr. Hellmuth Wachtel, Dr. Jim Seltzer, or Dr. John Burchard, all of whom have excellent academic credentials in biology and applied genetics as well as extensive experience in dog breeding and cynology, to confirm the validity of what I have said here.

We need to stop inbreeding Chinooks to death -- now! Would that we could backtrack twenty years and do things differently; that is unfortunately impossible. But for the sake of our own beloved animals' health and welfare, for the sake of every new Chinook litter that will be born in the next decade, action should be taken now. There is only one way to turn this kind of entrenched inbreeding around, and that is as I have just described: develop more outcross lines and release them for unrestricted breeding without backcrossing them to the point that no significant new gene inflow remains. As things now stand, the inbreeding will only increase inexorably. All that is needed to inbreed the Chinook to death is to do nothing. The present system can have but one outcome -- the death of the breed.

People seem to be worried about the Chinook Club of America and the American Kennel Club, as though if we were to disturb them it might somehow be the end of everything. I would suggest that they may well BE the end of everything if we do nothing and fail to make our own brave effort to solve the inbreeding problem. It may well be that no one in either CCA or AKC is capable of recognising the truth of what I have stated here. But those people in the end are not responsible for the Chinook as a breed, not yet anyway. COA is, at least for the present, and if the TRUE "parent club" of the breed sits on its hands for fear of AKC, that could seal the fate of Arthur Walden's tawny sleddogs. I hope that does not happen.


Post Scriptum - July 2011

The above commentary was written ten months ago. It seems appropriate to bring my thoughts up to date before closing and finally publicly posting this web page. In all honesty I have to say that my confidence in the future, the North American economy, the future of the Chinook breed, and the COA have all declined considerably during the past year.

In summer of 2010 three influential COA breeders viewed Tullibardine Duke Ellington, Darka of Seppala, Atholl Charles Edward Stewart, Atholl Lady Macbeth, Atholl Flora Macdonald, and Atholl Black Agnes Randolph. At that time, all three persons expressed great enthusiasm for the crossbred litter as Chinooks and as a potential Chinook genetic resource. It then seemed likely that we might have some help and support should we make an attempt to enroll the Duke/Darka litter in the COA Chinook Crossbreeding Programme

However, more recently each of those three individuals has separately given us to understand that it would probably not be worth our while to invest any money and effort into attempting to qualify Darka of Seppala as a Dog Zero and Tullibardine Duke Ellington as her mate. Of course nothing definite or specific was put in writing, but we were nonetheless left with the impression that to attempt to gain Chinook Cross status for the above Atholl litter would be a waste of time and would not be welcomed by COA.

Over the past year we have noted a quickening in the tempo of upgrades from Chinook Cross to "purebred" status. It seems likely that people are rushing to qualify their dogs as purebred ahead of full AKC breed recognition, and that the Cross programme will then become even more of a dead letter than it is at present.

Personally I no longer hold much hope for the future of the Chinook breed as such. The existing genetic problems are steadily increasing, while the will to remedy the situation by effective means (rather than relying on still more screening and selection within an already depauperate gene pool) just does not seem to exist in either camp, COA/UKC or CCA/AKC. So if ever there was a good time to retire from active Chinook breeder status, that time is probably the present. It seems a shame to see useful bloodlines coming to an end -- particularly diverse, quality lines like the Duke/Darka litter -- but if the desire of breed clubs and their members is only to produce more and more "purebred" Chinooks, sicker and sicker dogs, then it probably doesn't matter. If that's truly the case, then the situation is hopeless and breeding Chinooks is no longer a humane enterprise.