The Problem | The Legacy of Sir Gay | Looking for Diversity | What Remains to be Done?
What the Future HoldsThe Problem
We first started looking at Standard Poodle (SP) genealogies in search of an explanation for the prevalence of several problems that are either genetic in origin, or have a substantial hereditary component. This is not a problem unique to poodles, and is often due to inbreeding - the result of crosses between close relatives such as cousins. Inbreeding often occurs when a breeder relies too heavily on dogs from his/her own kennel, or the total population is too small.
However, a random sample taken from 1992 revealed that, in most 5-generation SP pedigrees, there were few ancestors shared by both the sire and dam. The average inbreeding coefficient was on the order of 3%, which is lower than that published for several other breeds.
Genetic diseases should not reach high levels in a population. Normally, one would expect them to be selected against, either by nature (the affected animals die), or by responsible breeders who would not use them in a breeding program. Unless individuals carrying these genes are inadvertently favored in a breeding program, selection should act to decrease their frequency. Unfortunately, there are several reasons why this may not happen:
Heterozygous advantage
The unwanted gene is selected for because the heterozygous condition confers selective advantage. A human example is the gene for sickle cell anemia, which increases the resistance of heterozygous individuals to malaria.
In a SP, one could be favoring bloat by selecting for a particular conformation that appears "attractive", or for hip dysplasia because a young dog only marginally affected seems to have better movement. Furthermore, though these problems may result in crippling or death as the dog ages, they may not be especially evident until the affected animal has been bred a number of times. Should such an individual become a "top producer", the problem will be passed to a disproportionate number in the next generation.
Too small a founding population
An unwanted gene for a genetic disease may be found only rarely in a large population. However, most individuals carry several such genes. Therefore, if a breed is based on a handful of founders, the particular problems that they carry will appear at a much higher than usual frequency among their descendants.
This does not seem likely to have occurred in the poodle. It is not a recent breed, and both the recorded pedigrees and written descriptions indicate considerable diversity. The North American SP heritage originates in at least 6 distinct lines (3 white and 3 dark) of Belgian, English, French, German and Swiss origin. The genealogical records indicate that, while the number imported from each line varied, the total number should have been more than adequate.
A genetic bottleneck occurs
The population undergoes a severe constriction in size due to disease, war, lack of popularity or natural catastrophy. In effect, there are no longer sufficient individuals to carry the full range of genetic diversity originally present, and genes may be lost accidentally.
The story of the modern American Standard Poodle really begins in the early 1930s, when the Blakeen and Carillon kennels imported blacks of both English and Swiss-German origin, and Swiss whites. Carillon bred mainly blacks. Blakeen bred all colors, a tradition continued by Bel Tor and Puttencove. At no time since the founding of this population do any of these factors appear to have had sufficient impact to produce a genetic bottleneck. Though some reduction occurred during WWII, the major impact appears to have been on the number of new Champions. Individuals representing the major founding lines that made it to North America before the war are readily identified in the fifties.
The Legacy of Sir Gay
Our first attempt to identify representatives of these founding lines in the 1990s relied on tracing the male "tail lines" (the father of the father of the father...). The first 30 pedigrees all traced back to Annsown Sir Gay, a black born in Connecticut in 1949. As Sir Gay was a descendent of the black German-Swiss Anderl line, we began to concentrate only on whites, but it wasn't until the 13th white that we found a current dog with a different tail line. (This one was a surprise in its own way, as it was a silver line imported from England to Canada in the 1960s.)
The first question to be answered was whether this was an accurate picture of the population as a whole, or an artifact of following only the male line. This requires a much more thorough analysis of pedigrees and the population as a whole, and is far from complete. However, we have now analyzed 18 black Standards in depth, and the same two individuals - Annsown Gay Knight (a son of Sir Gay, born in 1955) and Wycliffe Jacqueline (born 1954; all-time top-producing Standard bitch) - make the largest contribution to the genetic heritage of every one. On the average, these two account for a remarkable 55% of the total. Two other sons of Sir Gay, Bel Tor Morceau Choisi and Carillon Dilemma, and a granddaughter, Sedbergh Mitzi, each account for an additional 5%. This means that, in effect, breeding any two black Standard Poodles is like breeding first cousins (or closer), even though no ancestors may be duplicated in the first four or five generations. By the time you get to the 13th generation, Gay Knight or Jacqueline may appear several hundred times. Inbreeding coefficients calculated on a five generation pedigree are essentially worthless, and specialized software is required to extend these calculations to 10 or 15 generations.
This represents an "artificial" bottleneck, created by the popularity of the Wycliffe and closely- related dogs. There were a total of 667 new AKC champions in the 1950s, and they did not die from some mysterious epidemic before they could be bred. However, Wycliffe dogs sired 1/4 of the champions born in the 1960s, and when dogs of other heritage were not used, their genes were lost to the population.
At the extreme lie a few individuals who owe 100% of their heritage to 5 ancestors, all used by the Wycliffe kennel. Their inbreeding coefficients run as high as 74%! For the others, whose ancestry falls partially outside Wycliffe, a complete analysis is dependent on identifying "founders" in the other major kennels. (A founder is an individual living at the time of the bottleneck who is not an ancestor or descendent of another founders.)
(See also The Influence of Wycliffe on the Black Standard Poodle)
Looking for Diversity
Having discovered that a recent white derived from a silver line appeared to have somewhat different heritage, we started looking for white dogs with silver and blue ancestors. (When a black is bred to a white and one gets blue puppies, the silver gene is involved.) We have analyzed 10 recent white or silver Standards. Two owe about 60% of their heritage to the Wycliffe founders, and cannot be regarded as significantly different from the average black. Most of the others have a substantial Wycliffe contribution but, at around 35%, it is half that of the blacks.
What remains to be done?
- Complete the analysis of major kennels having significant "presence" at the time of the apparent bottleneck.
The objective is to establish what portion of the heritage of any current poodle is shared. Is it greater than the 68% already identified? Obviously, if most of this remaining fraction is "unique", the population, as a whole, carries more diversity than if it is not.
- Analyze a larger number of black poodles
We would like to have a representative sample of AKC and CKC champions, dogs holding obedience and other working/performance titles, and untitled dogs. The objective is to see whether the entire North American population is homogeneous, or whether dogs bred with different objectives have formed subpopulations. (The results of this analysis will determine how we approach the minority colors.)
- Analyze the minority colors
As we know the proportion of the different colors in the population as a whole, we will then be able to estimate the diversity in the total population.
What the Future Holds
By identifying individuals that owe a significant portion of their heritage to ancestors outside the "mainstream", concerned breeders can plan crosses that pair individuals who are not as closely related as they might otherwise be. This may reduce the risk of inheriting the same bad gene from both parents.
By encouraging their use, we may be able to preserve remnants of older lines whose genes will otherwise be lost. The key to long term survival of the breed is to reestablish a gene pool with greater diversity so that each problem is carried by a much smaller proportion of the population.
Regular reports of our progress toward understanding the genetic structure of the Poodle population will be appearing on this site and in The V.I.P. Bulletin, published bimonthly by Versatility In Poodles.
Individual owners and breeders can help by sending pedigrees and collateral data, thus enabling us to form a better picture of the entire population. The research database was started by an American genealogist from AKC stud books. It is believed to include all AKC show champions back to 1929, most English and Australian champions, and many others. However, it does not include all CKC champions, or many others with acceptable conformation that have achieved recognition in other areas.
If you are willing to help, or would like more information, contact:
Dr. John B. Armstrong
The Diversity Project
Department of Biology
University of Ottawa
Ottawa, ON K1N 6N5, Canada
email: jbarm@uottawa.caPoodle Links
- Poodle Support Group
- Versatility In Poodles
- Poodle History Project
- The Poodle Museum
- Wonderful World of Poodles
This information was last updated on April 18, 1997
© John B. Armstrong
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