Archive | Genetic diversity

Game mechanics and animal fancies


As anyone who is friends with me or my family members on Facebook can probably tell, we play a lot of games at our house. The guys play ruthlessly, so we often joke that it’s not a real game unless someone cries. As the mother, and resident peacemaker, this has made me keenly aware of the concept of game mechanics. Those are the factors that make a game hard enough to be a challenge without being needlessly frustrating.

The card above comes from the French game Milles Bornes. It was a game I played with my siblings as a child, but somehow I missed the fact that the game had terribly bad mechanics. Certainly they do not work for my own family! That’s because this card unbalances the game. It did not take long for my children to figure out that once you have this card, it is almost impossible to lose. So someone finally draws the Right of Way, and someone else cries. Whenever someone suggests this game, most of us just groan because we know what is coming. What we need to do is experiment with altering the rules to make the game more fair, but somehow we always end up using the traditional rules.

I have often wondered why people in the animal fancies – that is, the hobbies devoted to breeding and showing purebred animals – don’t question their own game mechanics.

I suspect that is because many are unaware that the “rules” the activity has are exactly that: game mechanics. Why do the spots have to be just so big, and just so regularly spaced? Why can the tail end be white but not the feet? Why does it matter if the white collar circles all the way around the neck? Do any of those things actually matter to the well-being of the animal? Do they contribute to his ability to do the job he was originally bred to do? Are they tied to an agreeable temperament?

The answer is usually no. What they are about is game mechanics. The game, which in this case is competitive exhibitions of animals, has to be difficult enough to keep the interest of the participants. By adding color requirements to the list of desirable traits, the game is made more challenging. The more specific the requirements, the greater the challenge.

Breeding to a standard is already a challenge. The head on a Collie, which is one of the most important aspects of type in that breed, is required to have very specific angles. The parameters of this are outlined in detail in the breed’s illustrated standard. (The Collie Club of America has one of the most instructive standards in this regard.)  Unfortunately the desired head shape fits between two norms that the canine skull structures tend to take.

CollieCompare copy

The middle image is the ideal Collie head. The heads to either side represent the extremes, with the “Borzoi” type roman nose to the left and the “Farm Collie” dished face to the right. One of the challenges to breeding Collies is getting that elusive middle image. If you are breeding Collies, that is part of the game.

For some animals color and markings are part of the challenge. Yet color is different in one significant way. Compared to something like the angles of the skull, color is pretty easy to predict. Take Boxer markings, for example. Boxer breeders like flashy white markings like the ones seen on these dogs.


In most breeds of dog, this kind of pattern where the legs, belly and collar are white is thought to be recessive. It is most often called Irish spotting, though in Danes it is called Mantle. Boxers are different in that while the white occurs in much the same areas, the pattern is incompletely dominant. Dogs with one copy look like the pair above. Dogs with two copies look like this.


Puppies like this are called White Boxers or Check Boxers (if there are significant patches of color inside the white). White and check Boxers cannot be shown. This is the American Boxer Club position on white Boxers.

ABC policy strictly forbids registering white Boxers with the American Kennel Club, as well as selling white Boxers or breeding white Boxers. The ABC also requires that white puppies not be included in the count on the AKC litter application form. The ABC has never condoned or encouraged the culling of white puppies.

Yet breeders know this is what they will get. It is entirely predictable. If breeders are using white trimmed dogs in their breeding programs (and trends that I have observed suggest that most are), then this is the fate of  25% of all Boxer puppies born in show breeding programs. If the average litter is eight puppies, then each breeding produces two “waste” puppies right out of the gate, before any other aspects of quality are going to be assessed. What if the hoped-for improved quality for that litter happened to fall to one of those white puppies? The parent registry requires that you not even retain the dog for breeding. That makes eight “waste” puppies – and another two additional ones that will be white the next time.

There are worse situations where the color requirements produce a lot of predictable culls. Harlequin Danes are the result of the merle gene and a modifier that strips away the silver color so that the background is white. Merle without Harlequin is not an acceptable color, but Harlequin is a homozygous lethal so all living Harlequins are heterozygotes. That means half the litter will not have the harlequin modifier and will be merle – a color not permitted. It gets even more complicated because merle causes defects (and the wrong coloring) in its homozygous form. That means ethical breeders will be crossing to non-merles, so only half of the puppies will inherit merle. If the breeder is lucky, most of those will be the ones that get the harlequin gene, too, but that’s not a given. To stack the deck even further, the Harlequin gene is “leaky” so undesirable merle patches sometimes show through. The potential for a lot of “pet quality” colors is pretty high.

Those are the rules of the game as it is currently played. Many of these rules date back to the founding of registries. Yes, that makes the limitations traditional, but it also needs to be understood that the men (and it was largely men) making those rules lived in a different world, and looked at animals differently. How easy is it to place a half-dozen off-colored puppies that will one day grow to giant size today, when many families cannot even fit a regular dog into their lifestyle? And are we comfortable with a built-in percentage of “waste” in a world where perfectly healthy animals go begging for a home?

Games only work if there is a challenge. The downside to this kind of challenge is that, depending on how hard it is to get it right, you are going to have a certain amount of misses. With animals, that means offspring that have no future within the fancy. Obviously we all expect fanciers to behave responsibly in regards to the excess animals they produce, and many breeders go above and beyond in this regard. But still the structure of the game ensures that some portion of the animals created will not have a place. When those undesirable animals are the sure outcome of something that is desired, then maybe it is time to reassess the rules of the game in light of the world as it is now, and our modern understanding of animals as fellow beings rather than objects. We do have the power to write new rules.

Update: It has been brought to my attention that the American Boxer Club website page on white Boxers does not have current information. In 2004 it was voted to amend the Breeder’s Code of Ethics so that breeders could offer Limited Registration papers and recoup medical expenses when placing white puppies. It should also be noted that until 1985 the Code of Ethics did not allow for placement of white puppies, which meant that while the statement that culling was never encouraged is technically true, the only action officially allowed was that the breeder keep every white puppy born.

(Boxer photos from Wikimedia Commons, Collie illustrations from the CCoA Breed Standard.)

Continue Reading

More on closed registries

(Photo by Tereza Huclova, provided for the upcoming book by the Kladruber Stud)

Several people have commented about the convex profiles of the Kladrubers and how they are visible even in the foals. I thought I would share this photo since it illustrates the characteristic so well. The extent of the arch varies from horse to horse, but it does seem especially pronounced in the black horses. As someone who grew up around Walking Horses when this type of head was not uncommon, I find it very appealing.

I also received a number of responses about genetic diversity and closed registries, so I thought I would include some links and reference material for those with an interest in the subject.

As Sarah Minkiewicz-Breunig mentioned in her comment, I like to recommend the book Bred for Perfection by Margaret Derry.

In the book Derry explores how culture, economics and the (then new) science of genetics shaped the world of stud books and animal breeding. She doesn’t shy away from the less appealing aspects of that history – namely the close association with eugenics and social Darwinism – but she does so in a less sensationalistic way than many.

There are also a lot of resources from the dog world, in part because rigidly closed registries have long been the norm within that community. One of the best sites for articles is the Canine Diversity Project.  Among the particularly good ones are:

The Poodle and the Chocolate Cake by Dr. John Armstrong  (a really good overview of the problem)
The Price of Popularity: Popular Sires and Population Genetics by C. A. Sharp

The dog world has also had some interesting experiments with out-crossing. Closed registries are the norm for dogs, so these have all been very controversial. Among horses relatively few breeds maintain completely closed stud books, and fewer still are willing to pay for that choice in the way that dog breeders are.

The Backcross Project  (restoring proper uric acid levels to Dalmations)
Bobtailed Boxers – Part 1
Bobtailed Boxers – Part 2
Bobtailed Boxers – Part 3
Bobtailed Boxers – Part 4
Bobtailed Boxers – Part 5
Bobtailed Boxers – Part 6
(This series is worth reading just to see how quickly a breed reverts back to type when crossed on something really different, all extensively documented in pictures.)

There was also a great post on the Terrierman blog about the importance of provenance in selling breeds.
You Can Blame Garrison Keillor’s Grandfather
The Terrierman blog has a lot of great posts about genetic diversity, though that recommendation comes with two caveats: 1) the blog is routinely sprinkled with unrelated politics that readers might or might not appreciate and 2) temperance and diplomacy are not really the style there. (He also takes exception to the color merle, which of course is not going to fly around here at the House of Blue Dogs!) Still it is consistently an interesting read, and it is worth exploring the (vast) archives there.

There is also the BBC program “Pedigree Dogs Exposed” which one of the comments referenced. To say that the program ignited a firestorm would be an understatement, and in parts it is certainly sensationalized. Still, what it portrays is happening, and perhaps something like this was the only thing that would push people to look at the issues involved. Since it aired, studies have been conducted and some preliminary actions taken by the British Kennel Club.

So what does that have to do with horses? To some extent horses and horse breeds are in better shape because they have not had a uniform culture of closed registries. Still where there are small populations and closed breeding programs, the same kinds of issues arise. Here is a discussion of the Friesian stud book policy on genetic disorders. This statement shows the very real parallel with the problems facing the dog world:

For this reason, the KFPS believes that carrier stallions should remain in the breeding pool. In addition, it should also be possible to approve young carrier stallions on the condition that they possess extra qualities. If we do not implement this policy, 1 in 3 young stallions will automatically be rejected on the basis of DNA testing.

One in three. And what are they carrying? Dwarfism. Twelve of the current stud book stallions are carries of dwarfism. Hydrocephalic Foals. Sixteen of the current stud book stallions are carriers of hydrocephalism. The effective breeding population is too small, and the defective genes so widespread that culling the carriers would probably narrow the gene pool enough that new defects would appear. That is what has happened in many dog breeds. It is exactly what should concern horse breeders, particularly those working with rare breeds with limited populations.

That is why many rare horse breeds have actually relaxed their color restrictions. They may still have strongly worded preferences for colors or markings, but off-colored or mismarked horses are not automatically removed in many breeds. Ironically, one of the breeds that does do this is the Friesian, which does not allow chestnut carriers as breeding stallions. This puts the registry in the awful position of permitting damaging defects if the horse is otherwise “of good quality”, but banning something as inconsequential as color no matter how nice the horse. As with dog breeders, this kind of disconnect with simple animal welfare undermines the assertion that purebred breeders are guided by a higher standard of ethics than “backyard” breeders.

Were it possible to expand the gene pool, culling could be accomplished without raising the inbreeding level to still more dangerous levels. Even without outright culling of carriers, introducing an outcross line can dilute the overall incidence of the defective gene. The fact that there are animals breeders out there who will choose “purity of blood” over solving serious health issues is something that deserves more attention. It isn’t a particularly comfortable discussion for a lot of breeders, and it points to the need for a very different mindset when it comes to breed stewardship, but it is one I personally feel must happen.

Continue Reading

My Ulterior Motive


My original purpose for the upcoming set of books was the explore the history of color in the different breeds. I did have another motive, though. It is impossible to tell the story of those colors without also telling the story of the breeds themselves, so in many ways the books are as much about breed history as they are about color. The idea of “breed” – and the related concept of “purity” – is often misunderstood. As anyone unfortunate enough to get me started on that topic is aware, I believe this situation harms animals. Purity of blood should never trump health.

That brings me to the horses pictured in this post. In writing the books, I have reached out to various registries and breeders for photographs. One of the most generous responses came from the Kladruber Stud in the Czech Republic. Countless pictures filled my inbox, many far more suited to a coffee table picture book than my more modest project.


Kladrubers are the last of a type of horse once known as galakarossiers, or ceremonial carriage horses. Kladruby nad Labem where they are bred is among the oldest of the European royal studs, having been established in 1560. The farm is currently on the Tenative List for UNESCO’s World Cultural Monuments.


Like most of the old European carriage breeds, the Kladruber is endangered. In writing the books, one of my hopes was to raise awareness of some of these little-known breeds. But in the case of the Kladruber, they also serve as a model for intelligent preservation of rare animals. There are two “breeds” of Kladruber, the Old White and the Old Black breed. This is in keeping with how color stood in for breed in the sixteenth and seventeenth centuries, by the way. The black variety was almost destroyed in the early twentieth century, with only one surviving stallion line and four purebred mares. Those directing the restoration made the decision to outcross to maintain the genetic health of the population, while still preserving the historical baroque type.

This wasn’t actually as radical as it might seem to those used to equating breed with purity. It was how Kladrubers were originally bred. It is how horses have historically been bred the world over. It is my hope that by telling the stories of these different breeds, more people realize that diversity and not absolute purity is really what is traditional.

And if that opens some minds to the less common colors out there, that wouldn’t hurt my feelings either!

Continue Reading