Friday, July 29, 2016

Stonehenge: The Root of the Inbreeding Problem

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 African. 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 reeding 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!


Gina Spadafori said...

Great read, Patrick ... but I do wonder about the Thoroughbred racehorses. I wouldn't hold them up as a shining example, COI or no.

They all trace back to three sires, after all, and more recently, the very busy and recently pensioned stallion Storm Cat* has been blamed (among with other factors) for the trend towards horses who race rarely, only a few times at all and are prone to fatal breakdowns. (Compare the number of races Seabiscuit ran in -- 89 -- with any notable modern Thoroughbred.)

If I recall ... racing Greyhounds have high rates of osteasarcoma, which doesn't make them a model of health long-term, either.

*Interesting side-note: Unable to impregnant T-Bred mares at the high rate expected of his formerly $500,000 stud fee, Storm Cat was retired fairly recently ... but then reported offered for breeding to select Quarterhorse mares. It may be that (as in Standardbreds), Quarterhorses can be bred AI, not "live cover" as required in T-breds. (Curious, I just sent a note to someone I know who covers racing to ask about this.)

Heather Houlahan said...

Working German shepherd dogs, like greyhounds, have very low COIs.

Winning show GSDs frequently have very high COIs.

As is to be expected when a strain has been selected for very rapid change to a uniform cosmetic extreme that is a drastic departure from the previous physical structure. And when health and talent are either not considered or considered a hindrance.

It's a shame that the chicken breed that bears Sebright's name is so damned delicate and unthrifty -- not to mention, purely ornamental in character.

PBurns said...

With racing horses and racing dogs, it is SPEED that is the requirement, and there is no other metric. These are not pets, but performance animals graded on a single axis, whether for good or ill. What is interesting to me is that inbreeding seems to hurt performance. Walsh suggests the same is true for bird dogs -- at least if it goes on for too long.

Pets are evaluated on a slighty different axis -- longevity and health. I would argue that if you are looking for those two qualities, the bias is almost always towards the smaller dogs. Under 40 pounds and over 10 pounds solves a lot of problems, while over 70 pounds or under 8 pounds brings a lot of problems with it no matter what the breeding.

I will write a wee bit more on Walsh, and note where the registries began to close up in another post. Let's just say that it was not done at the very beginning!


3Laiki said...

This Walsh fellow conjures up images of a 'sporting Martha Stewart'! LOL

At age 25, Storm Cat was pensioned at Overbrook Farm last May. It is not surprising that his semen quality was diminishing at that age.

QH mares can be bred AI and resulting offspring registered.

In the UK and Ireland, Thoroughbreds are bred for National Hunt Racing (Steeplechasing). They are bigger, more powerful, heavier-boned horses that mature later. In the US, the money is in short distances on dirt tracks for very young horses. There is a rush to get a return on an investment and the 'design' of the modern TB has changed since the days of Seabiscuit. The average number of starts and the rate of breakdowns has undoubtedly changed too.

For reasons above, people seek 3-day event horses in England and Ireland and cross their TBs to warmbloods and drafts...the latter being common sturdy foxhunting mounts ;)

Seahorse said...

The Jockey Club, the registration entity for Thoroughbreds, will not register any foal that is a product of anything other than a live cover. Not even if it is live cover supplemented by AI. Given that DNA-typing that is required to ascertain the parentage of each foal, live cover is certainly an antiquated requirement, not to mention more dangerous to horses and human handlers. Many, MANY other registries allow AI as a routine (not to mention the cattle industry's use for decades) and somehow have escaped whatever dark fate The Jockey Club seems to fear. Given the dire state of horse racing in this country, The Jockey Club might consider sobering up and modernizing their thinking. Not that I believe there should be even more breeding in general when today's foal crops are so casually discarded, but perhaps fewer horses would break down if breeding outside of one's geographic location was more easily possible. As always, there are many questions and concerns about breeding animals, but one thing is for certain: The Jockey Club is WAY behind in their thinking on AI, and doing the breeders no favors.


alfmcmalf said...

Brilliant piece. And a note on that dog show of 1859 this "There were sixty dogs entered, about 1500supporters attended and three judges were appointed for each breed. Messrs J Jobling, T Robson and JH Walsh judged the pointer classes and found an overall winner in Mr Brailsford's liver and white dog by Lord Derby's Bang out of his Dora. Setters were judged by Messrs F Foulger, R Brailsford and J. H. Walsh who found a winner in Mr Jobling's Gordon setter, Dandy. The winner of the pointer classes was owned by one of the setter judges, and the winner of the setter classes was owned by one of the pointers judges!"

From "Dogs, dog Breeding and the Control of Inherited Disease in the Dog" by Sampson, Irving and Willis. Published by our esteemed KC here in the UK no less.

I presume this type of "ticket" swapping is discouraged nowadays.


PBurns said...

It's commonly said that all Thoroughbreds can be traced back to just three stallions, but it's not true. You see, three stallions get you no progency -- so it's three stallions and a lot of mares. Of course, it also wasn't three stallions. In fact, over 150 stallions of "oriental " origin make up the lineage of the Thoroughbred and hundreds of mares too.

The Thorougbred line is 400 years old now, and there's been a lot of infusion and drift. Saying that a modern horse can be traced back to one of the original sires, is a bit like me saying I am related to Prince Charles or the Aghan Khan; even if true, it does not tell us much. There is a reason we do not prohibit second cousins from marrying, and why moving much past a 5-generation pedigree is pretty pointless in terms of genetics.

As for Greyhounds, the osteosarcoma here is not related to inbreeding, but due to the fact that the genes for size and speed seem to be attached to the gene that causes bone cancer. Genes are often paired (or more!) and the genes for size and speed are problematic this way.

PipedreamFarm said...

Microsatellite diversity, pedigree relatedness and the contributions of founder lineages to thoroughbred horses
E. P. Cunningham1,
J. J. Dooley2,
R. K. Splan3 and
D. G. Bradley1

Animal Genetics Volume 32, Issue 6, pages 360–364, December 2001

The thoroughbred (TB) horse is one of the oldest breeds of domestic animals, with pedigree records spanning three centuries. Because the population is essentially closed, there is concern about loss of genetic variation. Here we report two parallel analyses. In the first, genetic variation in the current population is measured using data from 13 microsatellite loci in 211 horses with relationships calculated based on allele sharing. In the second analysis, pedigree information is used to calculate genetic relationships between animals based on shared ancestry. These two measures of relationship are compared and shown to be closely related. Together, they provide an estimate of the amount of genetic variation which existed in founder animals. This study confirms the narrow genetic base of the breed and provides comprehensive analysis of contributions of founder animals. Seventy-eight percent of alleles in the current population are derived from 30 founders, 27 of these male. Ten founder females account for 72% of maternal lineages, while one founder stallion is responsible for 95% of paternal lineages.

dp said...

Think of Eight Bells and her pedigree.

dp said...

Is that book still available? Sampson was a great asset of the KC. Hknew what he was talking about.

dp said...

First cousins are allowed in humans afaik.

PBurns said...

Less of a book than a monograph. About $20 here >>