This article I already sended earlier to the list, but here again.

With thanks to Anne who helped me translating it to English

Rise of Dog Breeds

People and dogs have been evolving for tens of thousands of years. But the pedigreed pure breed as we know them today has only been in existence since approximately 1850.

Over the centuries, people created the dog.  Insofar as man bred critically, selection directed use and performance conformation.  Natural selection took good care that dogs could survive in a particular surroundings.  A hairless dog wouldn’t get far as a sled hiker on the North Pole. 

Slowly groups of dogs evolved with a particular purpose – watchdogs, pulldogs, herdingdogs, huntingdogs - and by its purpose and surroundings, people determined its exterior' - body, size and coat. 

For the new generation, people selected parents out of the group of workingdogs which were locally available. Therefore, especially in free insulated regions, the dogs inside the group started to resemble eachother. This is a reflection of accessory, not a breeding goal in itself.  'Good' became was measured as being effective, beauty was not an ideal to strive for.

Relative to the developments over ten thousand years, 150 years is of course nothing.  Yet there have been radical changes in the last century of dogbreeding.  The keyword for this change is the closed studbook, which, over the last half the nineteenth century has turned dogbreeding upside down. 

The Closed studbook.

Around 1850, the first breeds were ‘recognized’ and the first dog exhibitions were organized.  People founded breedclubs and wrote breedstandards, especially for conformation, and breeds were registrated. And they decided to register the lineage of the dogs. 

In the course of the time, for every breed, a lineage record was opened: the studbook.  In exhibitions and competitions, the best specimens of a breed were indicated, and those dogs were registered as a basic-generation in the studbook.  After a while, nearly all studbooks were 'closed', in other words, no new dogs could be registered, only descendants of in the studbook registered dogs.  From that moment, dogs were only considered 'purebreeds' if they were born out studbook parents.  The entire breed must subsist to the end of its days with the genetic material that was registered at the moment the stud book was closed.

The transition of genetically more open groups to a hermetically closed group of purebred islands where no ‘strange blood’ was permitted, meant a turning point in the dogbreed.  Pure breeding became a breeding goal on itself

(((An anecdote in this context is regarding a Dutch breed already in the first generations of pedigreedogs (i.e., three generations registered ancestors) where there was a high degree of inbreeding, through successive combinations of brother/sister, mother/son, father/daughter breedings.  When a breedspecialist fromof the beginning was asked to tell a little more about these dogs, because this surely had to be very special dogs, why else were so many inbreed, he had to laugh heartily.  Yes, they were indeed special dogs, they had a very desired characteristic that in a certain sense was 'heredity' namely: they had a pedigree!  And only from two registered parents you could get pedigrees of the pups - that is why they were paired to eachother!))) 

From useful to beautiful

More or less simultaneously with the rise of the closed studbook, for many breeds the job disappeared where they actually were bred for. Not only because their jobs were going downhill or were taken over, but also because people discovered breeds in far countries and removed them their original living and working surroundings.  In a large city there is of course not much 'useful purpose’ for a Saluki, a Deerhound or a Tibetan terrier. 

The useful aspect of dogs started becoming secondary.  Where, before, usability was the most important selection criteria for the dogbreeding, now the emphasis became more and more on appearance, the exterior of the dogs.  ‘Preservation of the breed’, the goal of every breedclub, was therefore translated as main reason for dog breeding and breeding dogs who fits the ideal of the breedstandard. 

Shows were organized to select the best breeding stock out of the shown dogs. To breed further with the most beautiful dog could also be worked on as the second objective of each breedclub: ‘Improvement of the breed’.

For dogs that did not fit the exterior standard, it was felt that they would not better the breed; rather that their genes would only blur the breed.  People didn’t realize that the profit they made on the outside beauty, finally would pay the price in the health of the later generations. 

This is the genetic basis of the most breeds, breeds that were already were serious limited by the closing of the studbook,

The Changes, In Summary

Seen throughout the history of dog breeding and in a very short time span, there have been radical changes in the way dogs have been bred.

After ten thousands years of gradual development of the different breeds by which:

- Natural selection took care of vital dogs, which could maintain vigor in the local environment, human selection mainly concerned working ability,

- Similarity in type among different breeds was a hazard rather than the result of selection,

- Genetic exchange between groups of dogs was possible

The purebred dog in the past century has been put in a straitjacket by:

- The desired appearance being defined by a breedstandard, instead of by the environment and the work the dogs had been bred for.

- Human selection which is mainly judged on outside beauty

-  Limited genetic exchange

- The genetic basic. Because the closed studbook was already limited, the genetic base was even further limited especially in cases where only champions were producing off spring.

With today’s basic genetic understandings which became available in the twentieth century, we now understand that the disastrous scenario. Today, nobody would dare recommend such a breeding policy.  However with the knowledge of 1900, people believed that this was the way to not only to keep the purebreed, but they even thought they could improve it in this manner. 

Knowledge of heredity in the past century

It’s well known that various qualities in a dog are hereditary.  But the understanding of how heredity precisely works, and what all will come along with it, is still growing.  Since the 1920’s, science has made enormous progress.  Scientific knowledge from 1950 was applied successfully in agricultural animal breeding, the management of in the wildlife and zoo populations.  But it would take years before the new obtained knowledge would be applied to dog breeding.

People still worked with the understanding from before 1900; thus breeders during a large part of the past century worked with an antiquated heredity model.  People understood that genes were the carriers for hereditary qualities, but thought that there were only two variants of every gene: a good version, that was normal, and a bad version, which showed up now and then. The breeders barely realize that the limited number of ‘clearly affected individuals’ indicated that at the same time meant that the population carried a large number of unfindable and intangible ‘carriers’.

People thought that for every quality, every characteristic, every body part, every illness, there would be a clearly apparent responsible gene.  The dog was seen as a whole that was built up was from loose puzzle pieces.  To breed the ideal dog you only had to replace the poorer fitting puzzle piece with pieces which would fill the hole

From this point of view, it wasn’t such a big leap to think that it was above all a matter of patience to get a beautiful and healthy breed. By breeding out the ‘wrong’ genes carefully and to bind the good genes, it had to be possible to get the breed or the own line purebred, with only the desired qualities.

‘Good breeding’ meant therefore: to trace the bad genes and breed them out, tracing the good genes and keeping them.


Inbreeding and popular studs.
Inbreeding – the mating of 2 closely related dogs – appeared to be the indicated way to accomplish the ideal of purebreed. With combinations of father and daughter, grandfather and granddaughter, mother and son, brother and sister you certainly quickly see what kind of genes you have. The ‘bad’ genes come out by inbreeding and can be eliminated so that only the ‘good’ genes will stay.

What breeders call linebreeding, is nothing more than a mild form of inbreeding, with some lesser related animals. But the underlying thought is the same: mating of related animals allows for tracking unwanted genes and removing them while keeping the desired genes

Inbreeding in combination with strict selection – excluding dogs with unwanted characteristics and only breeding the best dogs – quickly led to the desired result, certainly in terms of external characteristics. However, inbreeding also had very nasty consequences which became clear much later. 

With inbreeding and linebreeding successful kennels arose, breeding a very recognizable dog who regularly produced champions. Because many of those champions were tightly inbreed on 1 or several ancestors, it was highly like that they were pure for their fine qualities; therefore only had ‘fine’ genes to pass to their descendants.

That fortell of success made that they also by other breeders outside their own line much were asked a lot.  Everybody surely were willingly to breed beautiful and good dogs.  Sometimes such top-studs were so popular, that almost all breeders in a breed used them for their bitches.  The same thing happened with the sons and grandsons of these dogs, generation on generation on generation.  Nearly all breeds know such popular studs whose genetic material is spread over the whole breed. 

This manner of breeding - inbreeding, strict selection on exterior characteristics and popular studs – has lead to a situation where, in many breeds, genes from this handful of ancestors dominate.  All dogs then go back to the same ancestors.  Thus the intended 'purebreed' was in large part realized. 

Yet, not all present purebred dogs are champions, or even 'beautiful' in terms of the breedstandard.  And further, not to mention their health.  What went wrong then? 

Recent views
In the past few decades, the more recent scientific understandings permeated into world of the purebred dog. The realities of the genetics appeared to be more complicated than people had supposed. 

- The black-white-performance of a gene in two variants, good and bad, turned out to be incorrect.  In natural populations, there are several variants in circulation for every gene.  A few of those are really defective, and leads to illness (or be even death).  But most variants of a gene are said in more or less to some extent, 'good', or rather 'suitable'. 

- Genes aren’t regulated in nature for no reason.  If genetic variety is missing, all sorts of problems will appear, and there is a higher risk for illness, less resistance, and the adaptability declines.  A population with little genetic variation is a vulnerable population. 

For the health and chance of survival of a breed it is of big importance that for every gene several variants remain in circulation.  That means at the level of the individual, the dog, it is important to have at least for the most genes two different (suitable) variants per genepair.  Where one variant perhaps is lacking, the others can compensate, and so on. 

- Further it appeared that not all genes split, but are passed on in packages from parent to child.  These packages can contain genes for various characteristics.  Selection on one characteristic brings along, unintended, as well as a selection on other characteristics, that is always heritable (in pairs). 

And, to make it more complicated: the one-on-one-relationship between genes and their qualities is not accurate.  Nearly all characteristics - all biological functions as immune system, digestion etc., but also bonestructere, character – are affected by multiple genes.  That makes it even more difficult to assess the total-effect for selection of one quality. 

With these understandings, we can already see a large part of the problems.

Purebred or vulnerable?
First of all, the method of exclusion of 'wrong' genes was not as effective as was previously accepted.  The really big defects will manifest themself, and they can be contained, by selection - human or natural – for the most part.  But do we also notice when the genes only works 85%?  Or for 60%?  Probably not, also not as a dog two of such ‘a little bit defect' genes has.  We think that dog is healthy, sure if possible flaws - a weak heart for instance, or a form of cancer – show up at later age.  And if it is a beautiful dog there is probably bred with this dog as well, maybe even bred with a lot of times. And so ‘the little defect’ genes are spread in the breed.

Secondly the breedingmethods of inbreeding, strict selection and especially the exuberant use of popular studs, leads irrevocably to a decrease of the genetic variation.  Every individual has two specimens of each gene.  The number of genes in a breed is limited if more and more dogs have genes inherited from the same ancestor, then it only can mean one thing: that must have cost other variants of these genes, which are disappeared or at least have been decreased.  With these methods, genetic diversity has been decreased drastically - except that variety appeared to be indispensable to keep a breed vital!  'Purebreed' gets a total other meaning this way: a purebreed dog is vulnerable.

Thirdly it later came to light that breeders didn’t know which characterize they were selecting on.  With every puzzle part, others parts were coming with them.  That beautiful long fur went perhaps together with a little weaker lungs.  That fine, typical head brought perhaps a less breedtypical behavior.  And with getting rid of those ugly long ears, also thrown in perhaps was a part of the genes for good hips out the window. 

From evil to worse.
No single dog is totally free of defective genes.  Every dog (and every man) has within its thousands of genes 6 to 8 that are harmful, even deadly.  As long as there is a healthy gene of the same genepair opposite of it, the dog is unaffected.  As long as a dog hasn’t a large number of descendants, the breed will not be affected by the genetic burden, because of the miniscule chance that both parents with precisely the same deviation will meet each other.  In a natural population there are hundreds, possible thousands of different defects, but each in such small numbers that the harmful effect is negligible. 

With our breeds it’s different because there are no natural populations. The closed studbook was the start of genetic restriction. Inbreeding, popular studs and intense selection on a limited amount of characteristics, lead to an enormous impoverishment of the genepool. It is likely that many bad genes were removed, but at the same time other deviations, passed on by a few top-studs, were wide spread. These are now appearing in high percentages of dogs. The chance that two parental animals carry the same sick gene, and produce unhealthy litters, has become much bigger. While fewer diseases occur in the breed population, there are more dogs that have those minor illnesses. This is how breed specific sickness arises.

Even with relatively simple genetic deviations, where only one gene is involved, it usually takes a few generations before they show up. As long as both parents pass on the deviation, we unsuspectingly breed with carriers, who pass on the sick gene to half of their descendants. Until there are so many carriers that two carriers are combined, and the first sick puppies are born. Even then people often don’t realise that it’s a problem that’s passed on. Only when there are more cases get known, does it become obvious that the breed has a genetic deviation, by then it’s too late. The dog that introduced the sickness three or four generations back, has had so many descendants who have been used to breed with, that carriers are now through the whole breed.

As soon as a new genetic sickness appears, new selection rounds start. Carriers are traced and often are not allowed for breeding. A few studs from “clean” lines suddenly are popular. Their extra popularity leads to further impoverishment of the genepool – and in time, leads to the next genetic deviation. And so it gets worse everytime.

 
A part of the problems seems to be the consequence of a disruption of the genetic balance of the population. But there are more problems, and they also can be the consequence of a disrupted balance.

Disrupted physical balance

Striving to produce a “better breed” the breedstandard started to live its own live. At a certain point in time, the standard was made as a description in ideal terms of the group of dogs that belonged to the specific breed. But the description became exaggerated. When, according the standard, “small” was good, then “smaller” was better. When the skull was supposed to be wide, then the widest skull was the best. A long back could be longer, a deep chest deeper, a short nose shorter and so on.

But a dog isn’t just a collection of separate parts that, according to the latest trend, can be changed or modified. The primary purpose of his external appearance is not to look good. That exterior is the design of a physical balance, the balance that makes a dog a functional whole. No loose parts, but a whole. Changing one of the parts affects the whole.

For a bookshelf of one meter, two supports are enough. If we have a longer shelf we need more supports, or thicker timber, otherwise it bends. Everyone understands that. With a dog it’s no different. But people have only recently begun to realize this.

There many examples of breeds where the human impulse to change have caused a loss all proportions. A chest that’s so deep and narrow, that the stomach has no room anymore. Bloat is the result. Skulls that are so wide, that they cannot pass through the birthcanal anymore – C-section.

Backs that are so long that the legs cannot keep up anymore – Hernia. Noses that are so short the dogs have problems breathing. Skulls that are so tiny the brains don’t have enough space and come out. The list seems almost endless.

Natural selection would probably find a way to get rid of these problems, but we did our utmost efforts to disable this selection.

Disabling natural selection

Hundreds of years ago nature had a big voice in selection: weak, sick, unadapted animals had almost no chance of reproducing. There was no medical care to keep sick animals alive, and a bitch that couldn’t give birth the natural way didn’t survive, the litter also died. It was hard for an individual, but very important for the population, because this way the harmful genes could not be spread around. The most extreme cases of careless breeding was in essence corrected this way.

But in the past fifty years, veterinarian care has improved. For all sorts of problems and deviations remedies were invented. Cure, surgery, suppress, correct –veterinarian care gives us a broad scale of resources to help dogs with poor genetic luggage. This is good solution for an individual, sometimes lifesaving. The same medical care also makes it possible that a “repaired” animal takes part in reproduction, and that is a life -threatening situation for the population.

The harmful genes can now be passed on to next generations and a part of the natural selection for vitality is disabled that way.

Another effect of ongoing veterinary knowledge is that breeders can escape the long-term impact of wrong choices. The impact of inbreeding – as decreased resistance, decreased fertility – can be temporarily set aside by inoculations, dieting, antibiotics, hormone replacement, artificial insemination. The impact of extreme exterior – bloat, hernia, heart failure, difficult deliveries – can often be repaired by surgery. Many diseases that are often the impact of using popular studs – epilepsy, glands, pancreas, digestion – can be treated with medicines.

Not only did we take the ability to correct from nature, we slowly became accustomed to the consequences. “This breed usually has two or three puppies.” “This breed usually lives for eight or nine years,” “They always give birth through c-section.” We stopped asking ourselves if it’s all normal this way – we just don’t know better.

Inbreeding depression

It is difficult to tell, because processes as reduced fertility, decreased life span, and decreased resistance sneak in. After two, three, four generations, there is almost no noticeable change, and most breeders do not notice the difference. But when we compare multiple generations we see there is a difference.

The symptoms of decreased vitality of a breed is called “inbreeding depression”. The term speaks for itself: inbreeding depression is the consequence of individuals within a population who are too closely related. There is not enough genetic diversity, which is what makes the population vulnerable.
The combination of the closed studbook, strong selection on conformation, inbreeding methods and the use of popular studs leads to an enormous genetic impoverishment in our dog breeding.  We are backsliding.

It’s no surprise that we are starting to see the symptoms of the inbreeding depression. Allergy complaints, susceptibility to infections, studs that don’t mate, bitches who do not conceive, weak labour, adaptation problems which expressed by nervousness and anxiety, reduction of lifespan – these are all expressions of a loss of vitality.

With the help of antibiotics, infection-reducers, artificial insemination, c-section and valium, life is not so bad, but the root of the problems isn’t solved this way.

How further?

Deviations are part of life, and are relatively normal. It’s an illusion to think that it is possible to breed only healthy dogs. Sick dogs will always be born, we cannot change anything about that. What we can do is try to decrease the number of deviations. That the deviations come in percentages in our breeds, while in other animal populations they are expressed in the thousands, is unnatural, especially unnatural is the way how we breed our dogs. We cannot accept that a big part of the purebred dogs have so many genetic flaws, that they don’t have a life without medicine. We have to do something about it, that is obvious.

A look back at 150 years purebred breeding doesn’t only give us an understanding of what went wrong, it also tells us where to look for solutions.

Today’s problems are the result of:

- exaggeration of conformation features which undermines the biological functioning of the dog.

- large scale spreading of genetic deviations of a few individuals, through which high percentages of dogs in a breed suffer breed specific genetic deviations (popular studs);

- breeding methods that limit genetic variation and lead to inbreeding depression (inbreed, strong selection, popular studs);

The problems especially arise on the breed level of the population, the breed. For structural solutions the problems will have to be dealt with on the breed level. No breeder can do that alone, it requires a joint effort from breeders, breed clubs, judges and owners.

Animal healthcare moves on an individual terrain, not on the terrain of the population. Animal healthcare offers – much needed – outcome for animals that need healthcare, but cannot stop the perpetuation of those animals, and that is the problem we should address. It means that we cannot expect animal healthcare to find structural solutions.

How do we deal with exaggeration of conformation features?

Where health and well-beings issues are a straight result of an extreme interpretation of the breed standard, the solution is clear. The beauty ideal must be adjusted, in exhibition as well as when choosing breeding dogs.

In the past, through focussed breeding, physical features have been exaggerated so far that they became dysfunctional. Then it must be possible now, to correct the created damage in the same way, by giving preference to animals with less extreme features.

How do we deal with breed specific genetic deviations?

It is more difficult to search for solutions for the massive scale of spreading genetic deviations. These are almost always recessive deviations which means that we can recognise the sick ones, but not always the carriers, the dogs that pass on the disease without being sick themselves. Excluding all “suspected” animals – sick dogs, carriers and their close family – would be disastrous for most of the breeds, because then a big part of the genepool disappears. And we have seen where that leads to.

For some of the genetic deviations. DNA or marker research exists in the mean time for a few breeds. With a DNA research dogs free of genetic diseases, the carriers and the sick dogs can be identified at young age, before they are used for breeding. When used wisely the use of DNA research can help us breed healthy dogs.

This is possible by not using parents that both have the chance to pass on the defect gene (sick dogs and carriers).

To be sure we have to consider that DNA and other health research can cause other problems. The temptation is to breed to healthy dogs only. The result of that is inevitable: further impoverishment of the genepool, and in the long term new genetic deviations. Because all dogs carry some bad genes, if not for the one decease, then for another.

With DNA and other research we can hopefully try to diminish some of the genetic deviations. But with this method, we fight the symptoms of a much more damaging process, the loss of genetic variation. The cause cannot be dealt with this way. The cause is in the use of popular studs. And as long as we keep continuing that process, it will never get better.

We will have to keep cleaning for a while. But let us at least use our heads and limit the amount of matings for every stud. That is the way to prevent genetic deviations of one individual to spread across the whole population. Spreading risk is the only way to fight the problem of breed specific deviations.. Super genes do not exist, never have and never will.

What do we do about inbreeding depression?

To take care of the problems that arises with loss of genetic variation, we will have to do much more.

Limiting the numbers of matings for a stud is a very important step, but not enough. The still present genetic variation within a population will have to be used optimally. That means that breeding animals must represent the breed in a wide ranging way. If we want to maintain genetic diversity, and want to halt upcoming inbreeding, it’s no use to split 30 matings over four litterbrothers. Then we have to find less related dogs, and maybe give up our champions for a moment. Our breeding regulations must be checked very critically, and mustn’t have unneeded rules. We have to learn to respect the middle class of good functioning dogs (so called good and very good dogs), because that’s where the genetic diversity is.

And when the need is so high that there isn’t any diversity anymore, the we may have to go back to where the problem started, open up the registers, and consider outcrossing to a close related breed.

Inbreeding depression really exists, with dogs and especially with purebreds. Maybe breeding clubs would do well by removing “breed improvement” from their goals.

Let’s start with “health improvement”. After that we might hope for “conservation”.

Yes but,

With my breed it’s different …

The above sure is simplified. Not all breeds are mainly show breeds, not all the breeds have the same amount of genetic deviations, not all breeds show signs of inbreeding depression, and by far not all the breeds have suffered from deforming beauty ideals.

Let’s not think too much. For working champions the genetic laws are exactly the same as for show champions, for polygenetic deviations the laws are the same as for simple passed on deceases.

The processes are the same for every breed. Some of the breeds are at the end of the road, others are half way. It changes nothing about the fact that the road ends.