Saturday, October 11, 2008

Time, Volume, or Concentration

- - Why line breeding works, and works so very well if done correctly.

There is a pervasive notion in the dog world, especially among some of the FCI countries and in some working dog circles, that canine in-breeding is inherently "bad" and should be avoided. However, in-depth study of numerous successful breeding programs makes it clear that this notion simply does not hold up to objective scrutiny.

Some working houndsmen are quick to argue that many working dog breeds have developed "just fine" into their various functional types without inbreeding. What they fail to realize is that animal husbandry has "dimensions", and fixing breed quality in any domain(s) - functional, conformational, temperamental, etc - only comes with significant investment into at least one of these "dimensions." These dimensions are breeding "Time," breeding "Volume," or breeding "Concentration."

Straight away it must be understood that all of these so-called breeding "dimensions" are simply different ways of achieving the same goal--evoking the desired traits, and then hopefully fixing them in your line (or the breed population) through what would commonly be referred to as "Depth of Pedigree."

These dimensions, then, are the proven paths to achieving that depth of pedigree.

TIME - One way to achieve positive selection is to budget copious amounts of time to the breeding program. This only works with multiple -human- generations. Let's be clear, we're talking about centuries, or more often, millennia... not decades. A classic example of this is the Short-haired Borzoi, or more properly, the "Hortaya Borzaya." A truly ancient breed selected purely for field-proven function over millennia.

The Hortaya is considered to have been perfected hundreds (if not thousands) of years ago. As a 'perfected' breed, contemporary Hortaya breeders are strongly discouraged from line breeding (in-breeding) because functional selection is forgone; modern matings are instead determined by minimal inbreeding coefficients.

For the Hortaya, this makes sense. In a breed so old - and so relatively small - a population in which only field-proven animals are even considered for breeding in the first place, determining final selections based on maximized genetic diversity is an advisable course. That said, it needs to be very clearly understood that the important "take home" from a study of the Hortaya is that the breeders do not want true and absolute uniformity of type.

Hortaya breeders expect, and indeed want, several different types within their single breed.

Beyond the fact that most other breeds' breeders strive for absolute uniformity (at least in theory), the problems in attempting to apply this approach to most other breed populations is many-fold.

Most breeds are not nearly so old, and are not therefore, considered -perfected-. That is to say, dedicated contemporary breeders believe that they can not only maintain breed type, but further refine or improve it with future generations.

There is also breeder Ego to consider. Most folks want to live to see the fruits of their efforts to contribute to the breed. Individual humans have a finite life span. This means that they aren't willing to pursue a course in which they are merely an insignificant link in the chain of the breed's history. They want to produce the best dog possible... in their lifetime. For these breeders, the option of investing in the Time dimension, is not an option.

The major dichotomy of the Time approach is that many breeds which modern fanciers might think of as old, are in all actuality relatively young. Even so, they may in fact be relatively "perfected.” The relatively perfected state did not occur due to time at all, but investment in the breed's Volume, or the breed's Concentration, not all that long ago. A fact which is all too easily neglected and/or forgotten by the modern breeder.

VOLUME - It was not all that long ago that successful kennels produced some phenomenal individual dogs. We all look back and admire these dogs, and the breeders that produced them. What we neglect to remember is that they were, in effect, playing the Law-of-Large-Numbers.

Basically it says that in any process with variable outcomes, -every- possible result will eventually be achieved at least once within the parameters of possible outcomes, if (and only if) the process is repeated often enough. We conveniently forget that the ‘Annie Clark's’ of the world used to maintain kennels with dozens, or quite often hundreds of dogs.

Now before the working dog aficionado looks down their nose at this approach as the exclusive purview of 1920's show dog breeder, don't be naive... or hypocritical.

Especially those of us who admire the "older" continental and British bird dog and hound breeds. Those Dukes, Lords, Squires and Kings who kept and bred these dogs did the exact same thing. Keeping entire staffs dedicated to nothing but overseeing and tending to their kennels and breeding programs which usually included hundreds of dogs on site, and in a few cases, thousands of dogs.

The Volume approach then solves the Ego problem; it -does- produce results, and does so inside of one human lifetime. But there are two problems with the Volume approach as well.

The first 'problem' inherent to pursuing the Volume approach, is that it necessitates mass culling. I don't suppose to stand in judgment, and I am rarely want to ever tell another man or woman what he/she should do with their own dog(s). There is a time and a place for culling. The individual must decide what that means for them... only they know what will or won't keep them from being able to sleep at night.

The other problem with the Volume approach is the lack of an end-game. Followed in its purest form, this approach simply produces hundreds of puppies a year, secure in the knowledge that perhaps one in a hundred will "turn out". This dog (if it's a d-o-g) will be added to the stud stable, and hopefully, each year, he too will produce one in one hundred that is as good as, or better than himself. The other 99 will wash out, and likely become fertilizer.

The solution to this problem is at some point, investing in the Concentration dimension. A Volume breeder can spend the first 20 years of their breeding career producing a proven stud stable, and then line breed based on those foundation dogs for the next 20 years. But if this is the answer, then it also begs the question: If you're going to end up pursuing Concentration half way through, why not simply start there to begin with?

CONCENTRATION – The best way to effect the best results in the shortest amount of time, using the least number of dogs, is intelligent investment in the Concentration dimension. Concentrating is line breeding, yes even in-breeding.

The thing that needs to be understood is that line breeding does not produce, nor does it create, deleterious genes. All dogs - all of them - have deleterious genes. Line breeding simply "shows you what you got".

Let's say you are a bird hunter and by accident or by design you have acquired a dog with truly exceptional pointing ability. You've never seen another dog as good as this one. This is what you know. What you don't know is that this dog is also a carrier for "Three Leg Syndrome" or "TLS"- a syndrome I randomly made up which causes the effected dog to be born with 3 legs instead of 4.

So, in subsequent generations you wisely decide to line breed on this dog. What will inevitably happen? You will produce some affected dogs. That is to say, you will produce some dogs with TLS. That's the bad news.

This is where everyone starts waving their arms and stomping around indignantly. "See!!! This is why line breeding is so awful!!!"

But is it really? What was the alternative? Out-cross this dog to a (presumably) high quality and completely unrelated bitch. Then what? Did the TLS gene go away? Not likely. What will more likely happen is that the TLS gene will continue to hide in the pedigree for untold generations to come. Showing it's ugly head in the odd (seemingly) random puppy, and the breeder has no real or truly confirmable grasp on where in the pedigree it is coming from.

But using the Concentration approach, as effected dogs are removed from the gene pool with each subsequent line bred generation, fewer and fewer TLS carriers are produced, and more and more genetically clear dogs are produced. I refer to this process as 'dilution' to extinction through concentration.

In the modern age, 'dilution' to extinction can also be achieved without line breeding, at least for any genetic disorder for which there is a reliable genetic marker test. However, while strides are being made every day, most genetic disorders still lack a marker test. And even for those disorders for which there is a reliable marker test, what about breed type and functional type and improvement? Even when we can "test out" of a certain disease, out-crossed breeding paradigms still force us to operate in the Time or Volume dimensions to achieve maintenance or gains in those areas.

And herein lay the true beauty of the Concentration approach.

While we were busy producing more and more TLS clear dogs, and fewer and fewer TLS carriers and effecteds, through intelligent selection, we were simultaneously able to "set" that foundation dog's type features which caused us to select him as a foundation dog in the first place.

This is to say, with each generation, we can, if we so choose, get closer and closer to producing a clone of this now long dead ideal dog - without the TLS!

But some will bristle at the term "clone". Clone is not a dirty word. In fact, line-bred near-clones exist in nature. Dr Belkin:
In wild animals, you can see specialization for function that has evolved without regard for appearance. People select dogs to be dual-purpose - they just can't help it. The Arabs have selected Salukis not only to be the best dogs for coursing and catching things, but also to be pretty. Nature doesn't work that way because nature doesn't care about pretty. If an animal comes out pretty, that just happens... I like to think of the cheetah as God's Greyhound. The cheetah can do many things better than any coursing dog can do them. The conformation of a cheetah may tell you something about what our Salukis should be like if we breed them to course hares. Cheetahs are interesting in that not too long ago the world population of them was reduced to a very small number, so genetically they are all practically clones. They are about the most line-bred mammal in the world, aside from what has been bred by man. It pleases me to think that God got what He wanted and line-bred it.

Lest anyone mistakenly thinks that this approach is embraced by bird dog breeders and gräoid hound breeders, but avoided by trail hound and tree hound breeders, think again. Richard McDuffie, an old time (and I do mean old) coon-hound aficionado from the Carolinas, and a name worth knowing, recalls the story of "Woodrow's Dogs": When Woodrow returned home in 1945 after 4 years of military service, he was without a coon dog. He saw an ad in the local newspaper, the FayetteVille Observer, for 2 coon dogs. One was a Black and Tan for $50, the other a Bluetick for $100. He asked why the difference in price. The owner answered that the Black and Tan would tree most coons but not all, but the Bluetick treed them all. Woodrow bought the Bluetick and found that the man had told him the truth. He said that the first 32 tracks struck ended at a tree, and he shot out a coon. It didn't matter how cold a track was. If the old dog opened on it, he would put a tree at the end, and a coon would be there. Woodrow bred the Bluetick to some female the first spring. All the pups turned into coon dogs. He started inbreeding and only kept pups that were bluetick in color. The [inbred] dogs he produced generation after generation retained the traits of the original dog.

But wait. This has been been the functional reality of top performing tree and trail hounds since long before the 1940's. I won't detail the points of the article here (perhaps in a future posting?) but I will strongly encourage any of you who consider yourselves to be SERIOUS students of treeing and/or trailing hounds to write to the UKC, and request a copy of the article by Guy Ormiston entitled "A Tree Is Known By It's Fruit", which appeared in the August 2005 Coonhound Bloodlines magazine (vol 32, #8) in which Mr Ormiston details how the:

- Turn of the century (1800) Irish Hounds
- Captain, the foundation stud of the Henry Hounds
- Foundation stock for many running breeds including the Goodmans, Hudspeth, Julys, Shaver, Spalding Norris, and Triggs.
- Foundation stock for many treeing breeds including the the Redbone, English, Treeing Walker and Bluetick Coonhounds.
- "Tennessee Lead", pillar of the running Walker breed
- The leading field trial hound of the 1860's
- Many top field trial hounds of the early 20th century
- The 1928 National Leafy Oak winner

were -all- the product of rather intense inbreeding.

Why would this be the case? Because, I maintain, whether the breeder is consciously aware of it or not, 90% of what breeding quality dogs is all about is the cloning (or near cloning) of the most exceptional dogs. The other 10% is about making minor improvements in health, temperament, conformation and working abilities along the way.

If we stop and consider, clones (or near clones) actually offer us incredible selection options for the future. But how can this be? A line of clones is the apex of a bottlenecked gene pool after all. Well, this would be true of an entire breed population. But in most breeds there is a large population, and a large breeder base, so individual breeders applying the Concentration approach to their individual lines does not bottleneck the gene pool if Popular Sire Effect is avoided between lines.

So each breeder becomes the steward of a Concentrated line. Each of these individual lines becomes a little genetic island of healthy, typey, near clones.

The biggest problem inherent to the Concentration approach is that sometimes the most careful, thoughtful, and intelligent breeder will eventually encounter a genetic cul-de-sac in their line. This cul-de-sac is created due to a genetic defect that despite the breeder's best efforts, simply can't seem to be avoided within the line. Cul-de-sac can also occasionally occur due to the influence of a phenomena known as "inbreeding depression." This is where various reproductive qualities - sex organ formation, milk production, general virility, etc can be adversely effected, usually at a point at which COI's are extremely high.

This can be managed in several ways. Careful "back-crossing", an especially attractive option in the age of frozen semen, or conventional out-crossing.

Conventional out-crossing actually presents the most risk to the established and productive high quality inbred line. The effect of a Concentration approach is to greatly increase general homozygosity, and by default this greatly increases the uniformity and predictability of the offspring. A breeder who has invested many years or even decades into producing a high quality Concentrated line, opens their line up to all the unknowns and uncertainty of the other genepool when doing an out-cross.

That is unless the other line to which they are outcrossing is also highly Concentrated (intensely line bred), well established, and producing quality of type and health. Provided both breeders are honest with each other about what known deleterious genetics exist in their respective lines, they will quickly and rather easily know if an out-cross between the two lines is advisable. With a single infusion of new but concentrated genetic material then, the Concentration approach can once again be applied to many subsequent generations before the previously unavoidable defect and/or inbreeding depression appears on the horizon again, if ever.

As a perfect example of this, we can again look to the breeding program of Woodrow, the old-time breeder of jam-up Blueticks:
...the only problem with his dogs was that they tended to be ill at the tree. To prevent this aggressiveness at the tree, he outcrossed on a string of hounds known as Kentucky Bobtails that were being hunted in the Fayetteville area at the time and were known to be mild-natured. The outcross on the Kentucky Bobtails seemed to solve the aggressiveness at the tree. [As of 2006] Woodrow still had 11 dogs, all bluetick in color, that traced back to his kennel for 60 years.

While numerous varying but recurring archetypes may be considered acceptable or even desirable in a few breeds like the Hortaya, the vast majority of breeds are plagued by these seemingly unshakable variances. Unshakable for everyone except those rare Master Breeders who truly understand that beginning with the highest quality foundation stock available and then investing in that stock in the Concentration dimension of their breeding program is the most advisable course.

How have the Master Breeders come to this conclusion? They have realized that what they are really doing is breeding towards a single conformational and/or functional ideal. When that epiphany is achieved, and the breeder learns to make peace with it, it becomes clear that producing clones is not a bad thing... producing the highest possible quality clones is actually the goal. These are the breeders who throughout history have achieved consistent and long term success.

Once in a great while, a dog appears which is so close to that ideal that it distinguishes itself from the rest of its kind. The 'average' breeder dilutes this dog's quality by breeding it to a worthy but unrelated dog. The Master Breeder concentrates on this dog's quality through line breeding.

Is it a coincidence that true Master Breeders, both historical and contemporary, both working and show - the likes of Dr. Henry, Col. Birdsong, Dr. Belkin, Patricia C. Trotter, Dr Claudia Orlandi, and Guy Ormiston - all advocate(d) a breeding paradigm that achieves depth of pedigree (that is to say depth of consistent quality of pedigree) by spending the requisite amount of focus and attention to the Time and Volume dimensions of breeding, but focusing the lion's share of their focus and effort on maximizing their investment in the Concentration dimension of their breeding programs? Coincidence?

Long term and consistent quality is not achieved through coincidence.