Wednesday, June 04, 2014

John Henry Walsh and the Roots of Inbreeding

If you tease through history books looking for the roots of the Kennel Club, you eventually come to The Field magazine, and John Henry Walsh.

The Field was founded in London 1853, and its target audience was "those who loved shooting, fishing, hunting and could sniff out a decent claret at 1,000 paces."

In short, The Field has never been a pure hunting magazine; it's always been about social status as well. Articles include not only gun reviews, but also tips on how to select a good butler, and suggestions on where you can get the best catered lunch served to you on a linen tablecloth while shooting big game in Africa. Though The Field is the oldest country sports publication in the world, its circulation remains a paltry 30,000.

It's no surprise to learn that a publication written for pedigree people was one of the driving forces behind the creation of pedigree dog shows. In fact, The Field can properly be described as the birth place of The Kennel Club, and one of its early editors -- a former surgeon by the name of John Henry Walsh -- can fairly be called its midwife.

Walsh put himself out as an expert on nearly everything from home economics to cooking recipes, from brewing beer and playing croquet to shoeing horses, and from building and firing guns, to breeding and judging dogs.

The All-England Croquet Club, from The Illustrated London News,
July 1870. In the foreground: Miss Walsh, Mr. and Mrs. J.H. Walsh

Often writing under the pen name "Stonehenge," Walsh took charge of The Field as the first Information Age was exploding under the advent of low-cost paper made from wood pulp and movable metal type.

By diving into older texts, and compiling, rewording, and adding a little bit of new information, Walsh was able to liberate a great deal of basic knowledge and disseminate it out to an eager public.

Consider some of the publications Walsh wrote or edited between 1849 and 1884:

  • The Provincial Medical and Surgical Journal (edited 1849–52)
  • The Greyhound, on the Art of Breeding, Rearing, and Training Greyhounds for Public Running, their Diseases and Treatment (1853)
  • Manual of British Rural Sports (1856)
  • A Manual of Domestic Medicine and Surgery (1858)
  • The English Cookery Book (edited, 1858)
  • The Shot-Gun and Sporting Rifle (1859)
  • The Dog in Health and Disease (1859)
  • The Horse in the Stable and in the Field (1861)
  • The Shot-gun and Sporting Rifle: And the Dogs, Ponies, Ferrets Used (1862)
  • Riding and Driving (1863)
  • Archery, Fencing, and Broadsword (edited, 1863)
  • Athletic Sports and Manly Exercises (edited, 1864)
  • Pedestrianism, Health and General Training (1866)
  • Dogs of the British Isles (1867)
  • A Table of Calculations for use with the Field Force Gauge for Testing Shot Guns (1882)
  • The Modern Sportsman's Gun and Rifle (1882-84 in two volumes)

Walsh had a burning passion for horses, dogs, guns, and all things outdoors, and it did not hurt at all that he had been trained as a surgeon, as this meant he had an appreciation for the scientific method and could talk with some expertise about physiological problems common to humans, dogs, and horses.

Walsh's publication on the Greyhound was one of the first breed-specific publications ever produced, and Walsh was also a judge at the first real dog show, held in Newscastle-on-Tyne, in 1859. Ironically, there were no Greyhounds at this dog show: just 60 Pointers and Setters, with one class for each breed.

The same year that the first formal dog show was held (sponsored by two shotgun vendors it should be said), Walsh produced one of the first "all breed" books on dogs -- a publication which lifted much of its information from earlier authors who, in turn, freely plagiarized from even earlier authors.

As a result of copying from so many older texts, Walsh's 1859 publication contains solid fact which nests cheek-to-jowl with vague descriptions, absurd assertions, obvious truths, glaring omissions, and confusing verbiage.

It should be noted that Walsh's 1859 book and the first formal dog show coincide with the publication of Charles Darwin's Origin of Species. This is not a complete accident.

As noted in Inbred Thinking, both Darwin's work and the first dogs shows sprang quite organically from the work of Robert Bakewell and the stock shows he promulgated in the last half of the 18th Century.

The grass over Bakewell's grave had not yet grown over before farmers with a flinty eye on the steak-and-eggs axis of production, began to raise questions.

Among the first was John Saunders Sebright.

Sebright was a chicken producer and a falconry enthusiast who also wrote one of the first books on animal husbandry, entitled, The Art of Improving the Breeds of Domestic Animals.

First published in 1809, Sebright's book argued that the inbreeding and line breeding of animals, which had been so necessary to create breeds, could actually destroy them if continued for too long.

Sebright, of course, was right, and it was not long before progressive farmers began quoting Sebright at every turn as an explanation for declining fecundity and productivity in flocks and herds.

It should be said that Sebright was not some minor figure in his day. Darwin himself quotes Sebright repeatedly in his correspondence and books, and Sebright's ideas on the dangers of inbreeding were frequently cited in American farming periodicals as early as 1825.

Sebright's warnings were also picked up by those trying to tease out an explanation for the decline of Europe's aristocracy. In an 1838 publication entitled: Intermarriage: or the Mode in Which and the Causes Why Beauty, Health and Intelligence Result from Certain Unions, and Deformity, Disease and Insanity From Others," Alexander Walker not only quotes Sebright while framing the eugenics debate to come, he pecifically notes the parallels between the breeding of farm stock, dogs, and humans.

In one paragraph, he quotes Delabere Blaine, who wrote the first serious veterinary text on dogs:

That is as true today as it was then!

Walsh, of course, had read Blaine and Sebright. He quotes each of them once, and parrots their conclusions on a number of matters.

That said, Walsh's work is so slap-dash and hedging that it is often hard to tease out a straight sentence. As a result, when the reader does find a declarative sentence or two, they tend to leap out and perhaps be overemphasized beyond what the author intended.

Such may be the case on page 175 of The Dog, in Health and Disease, in which Walsh clearly sets out a series of "principles of breeding," one of which is rather boldly stated:

Breeding in-and-in is not injurious to the dog, as may be proved both from theory and practice ; indeed it appears, on the contrary, to be very advantageous in many well-marked instances of the greyhound, which have of late years appeared in public.

Now there's a declarative sentence! "Breeding in-and-in (i.e. inbreeding) is not injurious to the dog!

And yet, on page 188, just 13 pages later, Walsh writes:

The questions relating to in-and-in breeding and crossing are of the greatest importance, each plan being strongly advocated by some people and by others as strenuously opposed. Like many other practices essentially good, in-breeding has been grossly abused; owners of a good kennel having become bigoted to their own strain, and, from keeping to it exclusively, having at length reduced their dogs to a state of idiocy and delicacy of constitution which has rendered them quite useless. Thus I have seen in the course of twenty years a most valuable breed of pointers, by a persistence in avoiding any cross, become so full of excitability that they were perpetually at "a false point," and backing one another at the same time without game near them, and, what is worse, they could not be stirred from their position.

Eh? What the hell is Walsh prattling on about now? If inbreeding is not injurious to the dog, then why is any corrective breeding action needed?

You mean inbreeding can wreck the working ability of a dog? Eh?

What happened to that nice unambiguous sentence about inbreeding not being injurious?

And what's with all this nonsense about "once in and twice out?" What evidence does Walsh have that this is a "cure" for the problem which, just 13 pages earlier, he had denied even existed?

No answer is forthcoming, I am afraid.

The good news is that we no long live in the age of schooners and candles, and we can put Walsh's breeding theories to the test by simply looking at modern data for the Greyhound, one of the oldest breeds on earth, and the type of dog which Walsh himself knew best and cited as his "proof" that inbreeding does no harm.

Are today's Kennel Club Geryhounds deeply inbred?

No. In fact they are not. A recent Imperial College study of 10 Kennel Club breeds found Greyhounds had the lowest Coefficient of Inbreeding among the breeds studied -- and this despite the relatively small number of Greyhounds on the Kennel Club's roles.

We find extremely inbred dogs in each breed except the Greyhound, and estimate an inbreeding effective population size between 40 and 80 for all but two breeds. For all but three breeds, more than 90% of unique genetic variants are lost over six generations, indicating a dramatic effect of breeding patterns on genetic diversity.

OK, but that's Kennel Club Greyhounds.

Surely a great deal of inbreeding is occurring at the race tracks where speed is everything, and ethics must take a back seat to cash bets placed on performance animals?

In fact, if we look at racing Greyhounds we find a meticulous tracking of of Coefficients of Inbreeding (COI).

And why?

Simple: Because the higher the COI on a racing Greyhound, the less likely it is to be a winner.

Most racing Greyhounds have very low COIs, same as most racing horses.

In fact, with both racing dogs and racing horses, Coefficients of Inbeeding are tracked with precision and turn up on racing indexes where high numbers are treated as a bad sign. As one horse racing index notes:

[The Coefficient of Inbreeding] can generally be ignored unless you see a number over 5.00%. In that case, there's almost too much inbreeding in the horse's pedigree. A number over 10% indicates too much inbreeding and generally such horses don't do well at the track.

Of course, this is not the kind of information that John Henry Walsh was sharing with his readers, was it?

No. And why not?

Well, for one thing, Sewall Wright's Coefficient of Inbreeding calculation had not yet been developed.

Nor had the era of dog shows and closed registries begun. Yes, some people like Blaine and Sebright urged caution when it came to inbreeding animals, but aren't naysayers always with us? Progress always demands that we ignore the naysayers!

And then, of course, there was the matter of class and commerce.

John Henry Walsh could see there was a ready market for books about dogs, and the more breeds there were, the more words that could be written, and the more books that could be sold.

This is the same intellectual engine that drives the Kennel Club train to this day. The Kennel Club has shattered every type of dog into scores of breeds not because the dogs do different work, but because with every breed comes more ribbons and more people to chase those ribbons.

The business of the Kennel Club is not dogs you see, it is ribbon chasing, and it is not the dog that is interested in the ribbon, but the owner.

Of course, John Henry Walsh and The Field magazine meant no harm. They were simply trying to make a little money and gain a little social position by catering to a growing public interested in field sports and dogs.

And, as far as I can tell, though Walsh was clearly one of the hands pushing the business of dog shows forward, he also had real questions about the idea and nature of canine registries, even as he took the Kennel Club's money and acted as their publisher.

More about that in a later post. Suffice it to say that closed registries did not begin in 1859!


Dan said...

Just one small point of order here, on the subject of racehorses. They are in fact decidedly in-bred.

Thoroughbred horses came about with an import of a number of male Arabian horses in the 1700s, of which three were the most prolific sires. Of these, 95% of all modern thoroughbred males trace their Y-chromosomes back to the Darley Arabian sire.

A rather larger number of mares were originally used, but once again mitochondrial DNA typing demonstrates that only a small number of lines remain, with a reasonable estimate being a minimum of 15 and a maximum of 20.

Now, granted there are a number of other inputs into this pool, and I dare say quite a few mistakes over the years, but all the same a gene pool like this makes for quite a bit of inbreeding.

Thoroughbreds are therefore known for flat feet and for being of decidedly dubious temperament. Those which trace their ancestry to Oriole are generally somewhat lunatic and pretty stupid, indeed stupidity is a common thoroughbred trait and much skill in training is working out how to teach an equine moron the necessary racing skill set without losing it somewhere along the way.

This is why horse racing has such a high drop-out rate; you have to get every little factor spot on and teach the right things at the right life stages, along with cramming enough nutrition down the beast's neck to ensure it grows strong and fit. You also have to know how and when to run a racehorse without permanently turning it off racing.

Unknown said...

I have been following your blog for some weeks and stifling the urges to comment. I have admired working Jack Russels since my teens. Bought a British book in the eighties on the subject and saw that beagle was a frequently used outcross to keep nose,hunt, and to keep temperaments social. Always thought it was a sensible approach to dog breeding. My early experiences with AKC included Irish Setters and Bull Terriers so you are preaching to the choir. I hunt 13 inch beagles for rabbit and am amazed by how tightly bred many hard-hunting beagles are. I wonder if frequent culling is behind the seeming lack of negative effect. Or, is it possible that a genetically "broad" breed like the beagle( composite of hounds and terriers initially and able to hide outcrosses easily because of similarity to many other breeds) makes them resistant to some of the negative effects of in-breeding. I am raising two beagle/feist crosses now and looking forward to the fall. I have become addicted to your blog.

PBurns said...

This is a common misunderstanding of what it means to be "inbred" and is a routine mistake made by those who insist on running 10- and 20-generation COIs.

There is a reason that real dog and horse men (who are looking for animals with **utility** and therefore are rarely associated with rare breeds) do not go down that road. While you may get a number out of the equation, the number is no longer very meaningful as it is no longer a useful gauge of potential inheritance problems.

I address the issue here in some detail >>

Suffice it to say that both ancient law and modern law have no problem with you marrying second cousins, but both are rather concerned if you decide to have children with your mother or sister!

What the ancients knew, and what the moderns do too, is that genes "wobble" quite a lot and that genetic drift and mutation are such that a mathematically aberrant (and observable) genetic load does not occur in LARGE populations where close relations are not being bred.

To put a point on it, and to bring it back to dogs, if we look at Golden Retrievers, where there are millions of animals, the fact that **in theory** all of these dogs can be traced back to a handful of dogs owned by some British fop called "Tweedmouth" (I could not make that name up) back in 1900 (at about the the time the KC was closing the registries), does not mean all the dogs are horribly inbreed and diseased. In fact, the number of Goldens is jaw-droppingly enormous, and as a consequence Golden Retrievers, while far from absent health issues, are about as healthy as mongrels -- the "base line" for genetic diversity. See >> for the data.

What is true for dogs is also true for horse.

John said...

Having a closed registry does not necessarily lead to high degrees of in-breeding. The registries for all racing Greyhounds and coursing Greyhounds worldwide are closed registries and, as you point out, racing Greyhounds are not highly inbred.

Donald McCaig said...

Dear Patrick,
When in London some years ago I sought 19th century KC materials (and skeletons) at their library as well as 19th century copies of the Field. Destroyed in the Blitz, alas. Donald

PBurns said...

I cannot speak to beagles, but among working Jack Russells, there is very little inbreeding, COIs are tyopically at under 2 percent. Earlier this week I looked at a pedigree for someone else, and it had one dog, three or four generations back, that had a COI that hit 12%, which is the highest I have ever seen. That dog came from a Welsh hunt and from someone who knows working dogs well, so it might have been he was doubling down for a purpose. That can happen if you are trying to "lock in" a look, but if you are simply trying to make sure the next litter has nose or drive (what you need for work), that's simply too easy to find without ever getting near an in-and-in cross.

A lot of dog breeders, however are lazy or poor or lazy and poor, and often never go outside their own kennels for breeding. A reader on this blog sent me to a web site out of Michigan peddling Patterdales, and there I found pedigrees all of two-kennels wide. Yow!

Unknown said...

Patrick, I so enjoy reading your articles and this one especially was such a great read. On the matter of how far back to go on a pedigree to get a meaningful COI value, some breeders firmly believe that it is important to go back as far as possible. They don’t think going back only 5 generations suffices. You approach the matter rather differently however, but what you say also makes a lot of sense. It’s good to hear your viewpoint! I’m hoping your right as not having to go back so far would simplify matters tremendously for breeders and also not restrict further our already limited choices.

The Swedish Kennel Club apparently only calculates COI’s to 5 generations and they boast of an average COI for their purebred dogs of below 2%. Lena Gillstedt from Sweden had this to say about it……. “In small breeds, or breeds bred like they were small with a lot of bottle-necks, popular sires etc or when a breed goes from being very inbred to lo COI-breeding their COI can rise a lot if you calculate on 10 gens instead of 5. But in breeds that have been bred with low COIs for some time or larger breeds with less damaged populations, the COIs doesn't rise very much even if you go back 10 gens instead of 5. And the reason our pop geneticists chose 5 generations (all Nordic countries open databases plus Germany calculate to 5 gens) is that they wanted breeders to focus on stopping line breeding. Because new inbreeding is more dangerous than old inbreeding as it creates NEW problems. With the old inbreeding we see in the breed specific genetic disorders and immunological problems of many breeds. But to avoid NEW problems from continued inbreeding, 5 gens is enough. Old inbreeding due to the fact that selection, recombination and mutation somewhat reduce the effects of the old inbreeding over time. But of course the only true way to improve genetic diversity is to bring in new genes, as in outcrossing. But it is interesting to see that at least the purebreds in Sweden nowadays are being bred with lo COIs. And in one of my breeds, a breed having major health issues all over the world, we have insurance statistics that shows the breed is not so much worse off in total compared to other breeds.”

Lena goes on to explain how the Swedish Kennel Club accomplished educating their breeders…
“The first step was for the KC to acknowledge that inbreeding is a problem. Once they had done that, they could use all information sources available to inform the members.
Lots of articles in their own press.....Classes and courses for breeders.... Since a decade back, every breeder can use the SKK website with open databases to calculate the COI before a mating is done, and if it is too high, there is a red stop sign! Every litter and every dog has the COI information next to it in the open records. There are also graphs on COIs of every breed. In the beginning of 2000 all breed clubs HAD to make a document of the breed where they took insurance statistics and stats from the open databases and made an evaluation of the breed’s health status, this also contained info on inbreeding. Every breed put a goal to not let the COI increase more than 5% over 5 generations, and now a decade later ,the COIs are even below that goal.”

I think the Swedish Kennel Club took small steps but made a giant leap. I hope if nothing else all our main canine registries will follow Sweden’s lead and educate and empower their member breeders to have the desire and ability to calculate COI to at least 5 generations.

PBurns said...

I always look at the lay public's use of population data with some amusement. I once opened an American Association for the Advance of Science's symposium on Census adjustment by noting that in 1980, the population of the U.S. was determined to be 226,545,805. Then I paused, and looked up, smiled, and noted that even with the very low **acknowledged** error of the Census, the only digit that was certainly true in that number was the first digit!

The same is true for other kinds of population data. For example, people think a total fertility rate of 2.2 is horrible because it means population growth, while a TFR of 1.9 is horrible because it means population shrinkage, but the folks who cluck such concerns actually do not know how a TFR is constructed or what it really means. In fact, 1.9 and 2.2 is just wobble around replacement level. A yawn.

The point is that just because you can generate a number with a great deal of precision does not mean that you know what it means or when and where it is meaningful.

With COIs, you may be able to get really big numbers if you go back 10 generation, but the numbers are meaningless. The fact that some people think a 10-generation number is meaningful simply means they do not understand the number.

To begin with, a COI is a mathematical construct -- a cohort approximation of relations used as a synthetic predictability of the potential disease load in a given dog or litter.

In fact, a COI tells you nothing about a dog's actual health, and it does not even given you a clue as to what problems might crop up.

What the Swedes are saying is that in a really large population there is a lot of wobble, but that even in a relatively small dog population (say 5,000 dogs), the stuff more than 5 generations back becomes much less important IN THE REAL WORLD and what is much more meaningful is your actual knowledge of the dogs (and the people) in that 5-generation pedigree. Why the people? Simple: At some point you cannot or will not know about the specific dog, but you can and will know about the specific people. Honest men and women who have hunted or worked their own dogs for 20 or 30 years, and who do not breed too often are solid gold. They are not doing things cavalierly. They do not smell of the lamp, but of the dog. And it is the dog that is always the expert.


Derek Crankshaft said...

Fascinating! :)