Alaskan sled dogs (often called huskies) are unique in that they are an ad hoc breed–some say type–bred to pull sleds over snow, sans registry, that still exist entirely outside of the conformation show system. This is an attractive breed for Border Collie folks who envision their ideal herding dog in much the same light: purpose bred and free from the fancy.
The implication is that Border Collies (at least those outside the AKC) are free from the unintentional corruptions of the closed registry system, the inbreeding and loss of genetic diversity, the popular sires, the kennel blindness. Perhaps even that Border Collies are a recent and ongoing amalgam of numerous breeds and strains of dog, a little of this and that to keep the gene pool fresh. Genetic research, however, shows that Border Collies are not like either of the two styles of Alaskan sled dog, and the comparison fails.
A 2010 study looked into Alaskan sled dog genetics and offers the following reason why they are an attractive dog for study and a wishful analogy for Border Collie elitists:
The Alaskan sled dog offers a rare opportunity to investigate the development of a dog breed based solely on performance, rather than appearance, thus setting the breed apart from most others. Several established breeds, many of which are recognized by the American Kennel Club (AKC), have been introduced into the sled dog population to enhance racing performance. We have used molecular methods to ascertain the constitutive breeds used to develop successful sled dog lines, and in doing so, determined the breed origins of specific performance-related behaviors.
The first revelation of the study is that even Alaskan sled dogs aren’t “bred like Alaskan sled dogs,” as there are two identifiable strains that are bred quite differently: Sprint Sled Dogs and Distance Sled Dogs.
We observe that the Alaskan sled dog has a unique molecular signature and that the genetic profile is sufficient for identifying dogs bred for sprint versus distance. When evaluating contributions of existing breeds we find that the Alaskan Malamute and Siberian Husky contributions are associated with enhanced endurance; Pointer and Saluki are associated with enhanced speed and the Anatolian Shepherd demonstrates a positive influence on work ethic.
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Sprint and distance sled dog racing are vastly different in terms of the distance traversed during a race and the speed at which this is accomplished. Long distance racing includes races of several hundred miles lasting multiple days, such as the Yukon Quest and Iditarod of over 1,000 miles in the subarctic winter. Sprint racing is more analogous to track and field with multiple competition events defined by the size of the dog team. The extreme differences in these racing styles, ranging from 30 miles in one day to 1,000 miles in less than ten days has lead to a divergence within the Alaskan sled dog population based on the essential physiological athletic attributes of endurance and/or speed as well as “work ethic,” which encapsulates an animals’ desire to perform.
It has been claimed by several sheeple that Border Collies are very much like Alaskan Huskies. Eileen Stein is the current President of the American Border Collie Association and has served on the Health & Genetics committee for several years, she should be in the know about BC genetics and she feels that the Alaskan Husky is an appropriate analogy for the Border Collie in how they are bred.
Eileen Stein – 9/22/2010 I think border collies ARE a breed like Alaskan Huskies. They are like Alaskan Huskies in that they are bred to a working standard rather than an appearance standard, and they are a breed rather than a type in that they have been bred to that working standard long enough that they almost always meet that workiing standard better than any other kind of dog. I don’t really understand why you think dogs must be bred for aesthetics, or must have no significant variation conformation-wise, in order to be a breed.
Alaskan Huskies don’t have a registry and border collies do — that’s the only significant difference. And probably that’s the only thing that has kept the AKC from going after them.
So how are Alaskan Sled Dogs bred? What do the numbers say?
First, Alaskan Sled dogs as a whole benefit from having two sub-populations under the same mental model of the breed. The Distance sub-population boasts a mean 2.9 alleles per locus and the Sprint sub-population has 3.1 alleles per locus and these alleles are mutually unique, as taken together the entire population has a mean alleles per locus of ~5.7. A combined set of 141 purebred populations, including Border Collies, showed only 2.6 alleles per locus.
Second, both Alaskan Sled dog types show a diversity in genotypes as well. The Observed Heterozygosity (Ho) for the Distance dogs was 55.9% and 61.1% for the Sprint dogs. Purebred dogs managed only 43%.
Third, the Alaskan Sled dog population shows a divergence in inbreeding. The F(IS) statistic is a type of inbreeding coefficient that ranges from -1 (highly outbred) to +1 (highly inbred). Negative values indicate recent outcrossing (a diversity of alleles that are trapped in a few individuals versus being evenly spread across the population) and if the assumptions of Hardy Weinberg equilibrium are met, it is expected that heterozygosity would increase with random mating.
Positive values indicate recent inbreeding (which can include close mating according to pedigree, non-random mating of like individuals, reduced genetic diversity due to population structure like subdivision or bottlenecks, etc.). Positive F(IS) values indicate that heterozygosity will decrease in the future, negative values indicate that future offspring will be more heterozygous.
As you can see from the above diagram, the Sprint Sled Dogs (purple) have an excess of heterozygosity representing active outcrossing and an influx of genetic material. The Distance Sled Dogs are more typical of other breeds, showing continued selection for conformity. Border Collies, unlike their relative the Collies and the Cardigan Corgis, have a positive F(IS) value indicating ongoing inbreeding.
Border Collies aren’t like either of the Sled Dog sub-populations. Even though the Distance dogs have higher F(IS) values, they are still highly heterozygous and have a greater abundance of allele diversity. In other words, Distance dogs are being pushed genetically towards homozygosity faster than the Border Collie is being pushed, but the Distance dogs are starting from a more diverse and less inbred position.
The Sprint dogs not only have a greater abundance of allele diversity and a greater level of Observed Heterozygosity, they are also being actively and continually outcrossed. This simply isn’t the case with Border Collies.
Border Collies have a virtually closed breeding pool of dogs that go back to a few hundred founding dogs a century ago. Their effective gene pool is now equivalent to the genome of only 8 dogs. The number and impact of new blood (typically in the form of Registration on Merit) is negligible. The contribution of other breeds (like Kelpie and Bearded Collie) is highly limited, mostly ancient (a century ago), and not ongoing. The last documented non-Border Collie to enter the gene pool is almost 30 years ago with one Bearded Collie (Turnbull’s Blue) ROM’d within the ISDS.
The last time a Husky was improved with fresh blood was probably yesterday.
The genetic analysis of the Sled Dogs indicated 21 domestic breeds that contributed to the Sled Dog breed with a population score of ≥ 0.3.
The 21 “related breeds” included the Alaskan Malamute and Siberian Husky, which were expected based on historical information, and the Pointer, which has recently and repetitively been bred into the population. The Samoyed, Chow Chow, and Akita also have historical roots as northern draft dogs. Other breeds included in the “related breeds” group were the Saluki, Afghan Hound, and Borzoi, which are well known for their speed, the Great Pyrenees and the Anatolian Shepherd, both of whom are northern climate guard dogs, and the Weimaraner, a hunting breed of shared ancestral heritage to the Pointer. Additional related breeds were the Japanese Chin, Shar-Pei, Shiba Inu, Shih Tzu, Pekingese, Lhasa Apso, Basenji, Tibetan Spaniel, and Tibetan Terrier, most of whom share an Asian heritage with the exception of the Basenji.
Thus, it is folly to suggest that the Border Collie gene pool and community breeding ethic is substantially similar to the culture of breeding working Huskies. This isn’t true. Huskies are an ad hoc landrace where registry politics and breed purity don’t matter. If you want to outcross your dog to an entirely different breed, you can, for any reason at any time without any ROM hoops to jump through and clearly racers do.
If Border Collies are substantially similar to this breeding ethic, find me one trial champion dog from any country that has a non-Border Collie in the last 3 generations. Heck, find me a dozen.
The interesting question is not so declarative, simple, and overstated as “Border Collies and Sled Dogs are both bred to a performance standard.” Yawn. You can make that statement without a comparison, so if we are to compare we have to ask more than just distilled philosophy and slogans. The question is HOW are they bred and what is the genomic situation in the breeds.
The answer to that question doesn’t provide an impressive comparison which elevates the Border Collie vis-a-vis the sled dog.
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Labrador retrievers, English pointers, and Llewellin setters are bred for trials. Their registries are not open, and they aren’t particularly genetically diverse, within their strains. Border collies are much more like those dogs than Alaskan huskies.
Another analogy would be FCI-registered West Siberian Laika since the RKF studbook is closed.
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I wonder if they would ROM a known mix? All the BC boards say that herding instinct is easy to lose if you don’t breed for it, but I don’t think that’s true. Certainly nobody has bred Shelties for herding instinct for a very long time (maybe EVER) but many show at least some instinct. If you had a BC bitch with strong instinct and bred her to, say, a whippet/staffy cross, and were able to prove the resulting offspring had the requisite ability to perform, would they ROM those dogs? Not an experiment I want to take on, but is someone considering something like that? You would think the advantage of a working registry would be to be able to bring in new blood, which obviously is not happening now.
I guess it’s a matter of what one sees as instinct. A good BC is fine tuned. But I’ve seen mongrels out of shelters move sheep around, cut them off, corner them, etc.
Rotties, rhodesian ridgebacks and kerry blue terriers have earned herding titles of some sort.
“Instinct” is a large topic.
Well, a “fine-tuned” BC has had a lot of training. Many, many breeds and mixes have instinct. I don’t know what the ROM-in requires–just wondering if a known mix could meet the requirements, would they allow that dog in, and allow it to be bred to “purebred” working dogs?
Aaaannndddd… I looked over the requirements to get a dog in through ROM, and you’d have to be pretty damn determined to do it–willing to spend a lot of money and time to prove a point.
Yes, and they could prevent you from getting the dog registered with just one abstaining vote from the board of directors.
Exactly. I’m willing to bet that sufficient “herding instinct” shows up all the time in mutt mixes and other breeds, but a) who would want to go through the bother of ROMing their Rotty, and b) I’d also bet that trial folks would HATE having a dog out on the field and breeding with their dogs if it didn’t look like a BC or similar.
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The point of a ‘breed’ for most people is predictability. A Rottie, as an example, is not, in most cases, a dog bred for herding, and definitely *not* a dog bred for herding trials. Picking a random dog with herding instinct, proving it, and then breeding it into the BC population would be, frankly, an exercise in futility, due to the predictability factor and how risk averse the breeder culture is. You would be unlikely to find people willing to breed to your dogs with the Rottie (or whatever) behind them. You would have a better chance with a less ‘risky’ dog, like a Beardie from working lines, or an English shepherd. Breeder culture, in the end, is what decides what a breed *is*, and what it *becomes.*
I am willing to bet that some of what you are seeing in the differences in homozygosity between sprint and distance huskies is down to differences in risk acceptance in their respective breeder ‘cultures.’ IOW, willingness to take a chance and breed anything that seems to have the desired qualities, versus playing it safe. An experimental nature, if you like. It is probably highly unlikely that the purebred dogs used in many of those experiments came from proven competition lines, as many of those people are thoroughly indoctrinated against allowing their stock to be used to create ‘mutts.’ A conservative culture. Which makes using such ‘unproven’ dogs a even higher ‘risk.’
(“I have no objection to lurchers or longdogs but I sure as hell wouldn’t sell a dog to a breeder or allow one of my stud dogs to be used by one,” is something I have seen many, many times with Saluki breeders. Part of that is the idea that the Saluki [or whatever purebred] is the ‘ultimate’ endurance running dog [or whatever] and therefore there is something inherently hinky about using one to make mixes, as *of course* mixes would be inferior.)
You do see some extreme divergence in philosophy even within a breed culture; there are certainly plenty of lurcher/longdog breeders who have formulas that they like (3/4 Deerhound, for example), or who absolutely would *never* breed to an unproven dog, and on the opposite end of the spectrum are those who are highly experimental and will try anything that seems interesting. I have seen longdogs used for hunting jacks and coyotes that looked like large, robust Greyhounds, *except* that they were saddle tan merles. The merle came from a catahoula and the saddle tan from a foxhound. This same breeder also experimented with Afghan hounds and Italian greyhounds.
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I know this wasn’t the exact point of your post, but when I checked out your H-W table I could not help but be surprised that collies and dobermans were in the negative.
Really????
I would have thought them to be sooo much less genetically diverse than that, whatever their total population numbers.
Yes, a shock. Now, you have to remember what that figure is measuring, and a more complete analysis needs to be done, but I have for more than two years been trying to collect an extensive list of observed heterozygosity results to pair with those deviation numbers.
You might think of the above chart as acceleration, the heterozygosity as velocity and the number of alleles per locus as distance away from 1.
You start with many alleles and you speed toward one. How homozygous you are grows as you accelerate.
So what were the breeds with some of the longest bars to the right? Scary looking rate of acceleration!
I have an old draft waiting for more data called “How inbred is your dog” that covers that in more detail. I’ve been trying to ferret out the Observed Heterozygosity numbers from the researchers for more than a year.
A new comment to this post prompted me to revisit, and now that I am aware of the prevalence of DCM in dobermans, the negative value for them in this chart has me even more curious about what outcrossing event occured.
You do have to remember what these data are saying. They’re not measuring Heterozygosity (Dobes could very well be highly highly inbred), they’re measuring the difference in what was expected from what was observed. In a way, it’s a lot easier to be a surprise if you’re dealing with a highly inbred breed already (like an F student getting Cs versus an A- student getting As). Getting observed Heterozygosity numbers for a wide number of breeds has been a task I’ve been working on for many moons.
Right. Not measuring overall heterozygosity. But as practically ALL purebreds well known to the public seem rather inbred, would it be fair then to say that this was RELATIVELY..uh, not so bad?
Not as bad as “expected?”
Hardly means there isn’t a long way to go and the recent outcrossing suggested might not have had an affect on whatever alleles are related to DCM ( as far as I know this info has not been definitively acertained).
But I wonder now, where and what exactly this result points to in the breed.
WAS there a recent outcrossing? Which dogs gave this result and why?
What else should I know Christopher? What might I not be seeing? 🙂
I don’t have an answer to your questions, you’d need a lot more information about the dogs than just the one number. But it’s undoubtedly a good thing as it means there is either active pressure or selection for greater heterozygosity and/or an outcross of some sort.
Either way this continued trend would lower the average homozygosity in the breed.
I guess that depends on what you are defining as “instinct.” If you mean sustained interest in livestock and an ability to use pressure to move them then many breeds can do that. My old Siberian could quietly move sheep around a round pen.
Theres more to a good herding dog than instinct, however. Its a combination of traits: biddability, confidence but not aggression (stand down the nasty ewe, but don’t eat her), light and fluid movement to not waste energy and handle huge outruns, stock sense (how close can I move without the sheep moving, enough obsession to stay with the job even when scared or tired but not so much they run the sheep into the fences, etc.
Getting all of those qualities in one dog can be tough.
I don’t know that much about mushing, but from a layman’s POV there are fewer fine tuned skills needed to be a good sled dog. I am sure someone will correct me if I am wrong.
Hi Chris, I was just wondering if you have a clearer, hi-res version of your H-W graph? When enlarged, I couldn’t read the abbreviations for the breeds. Or if you have one with the full breed listed out? This is something I’m really interested in having on hand.
I don’t have a larger photo, but I do have the raw data which will hopefully serve your needs:
Population f Breed
ACKR 0.117224 American Cocker Spaniel
AESK 0.077005 American Eskimo Dog
AFGH 0.053138 Afghan Hound
AHRT 0.032622 American Hairless Terrier
AIRT 0.088701 Airedale Terrier
AKIT 0.097219 Akita
AMAL 0.124276 Alaskan Malamute
AMST 0.086404 American Staffordshire Terrier
AMWS 0.224458 American Water Spaniel
ATOL 0.030853 Anatolian Shepherd
AUCD 0.080901 Australian Cattle Dog
AUSS 0.177875 Australian Shepherd
AUST 0.113363 Australian Terrier
BASS 0.097893 Basset Hound
BEAG 0.191412 Beagle
BEDT 0.344988 Bedlington Terrier
BELS 0.106362 Belgian Sheepdog
BERD 0.286425 Bearded Collie
BICH 0.090351 Bichon Frise
BLDH 0.049555 Bloodhound
BMAL 0.021066 Belgian Malinois
BMD 0.213088 Bernese Mountain Dog
BORD 0.017215 Border Collie
BORT 0.058169 Border Terrier
BORZ 0.042549 Borzoi
BOST 0.078869 Boston Terrier
BOUV 0.058605 Bouvier des Flandres
BOX 0.203778 Boxer
BRIA 0.051442 Briard
BRIT 0.018479 Brittany
BRUS 0.003066 Brussels Griffon
BSJI 0.129814 Basenji
BULD 0.113042 English Bulldog
BULM 0.102236 Bullmastiff
CAIR 0.114173 Cairn Terrier
CARD -0.05649 Cardigan Welsh Corgi
CCRT 0.273733 Curly Coated Retriever
CESK 0.218036 Cesky Terrier
CHBR 0.078677 Chesapeake Bay Retriever
CHIH 0.12681 Chihuahua Long Coat
CHIN 0.100481 Japanese Chin
CHOW 0.155405 Chow Chow
CKCS 0.185008 Cavalier King Charles Spaniel
CLSP 0.019274 Clumber Spaniel
COLL -0.01916 Collie Smooth
CRES 0.155039 Chinese Crested
DACH 0.073893 Dachshund Wire Haired
DALM 0.121302 Dalmatian
DANE 0.242629 Great Dane
DDMT 0.197501 Dandie Dinmont Terrier
DEER 0.029535 Deerhound
DOBP -0.01482 Dobermann Pinscher
ECKR 0.080039 English Cocker Spaniel
ESET 0.149783 English Setter
ESSP 0.007521 English Springer Spaniel
FBUL 0.224551 French Bulldog
FCR 0.110334 Flat-coated Retriever
FIEL 0.053266 Field Spaniel
GBGV -0.02908 Basset Griffon Vendeen Grand
GLEN 0.04904 Glen of Imaal Terrier
GOLD 0.059375 Golden Retriever
GORD 0.140875 Gordon Setter
GPIN 0.062108 German Pinscher
GPYR 0.00415 Pyrenean Mountain Dog
GREY 0.242856 Greyhound
GSD 0.09771 German Shepherd Dog
GSHP 0.352865 German Shorthaired Pointer
GSMD 0.133979 Greater Swiss Mountain Dog
GSNZ 0.137121 Schnauzer Giant
HAVA -0.04366 Havanese
HUSK 0.120417 Siberian Husky
IBIZ 0.137128 Ibizan Hound
IRSE 0.029264 Irish Setter
IRTR 0.122333 Irish Terrier
ITGY 0.20339 Italian Greyhound
IWOF 0.189741 Irish Wolfhound
IWSP 0.026804 Irish Water Spaniel
JACK 0.029031 Jack Russell Terrier
KEES 0.098407 Keeshond
KERY 0.117097 Kerry Blue Terrier
KOMO 0.020065 Komondor
KUVZ 0.077393 Kuvasz
LAB 0.1564 Labrador Retriever
LANC 0.005155 Lancashire Heeler
LEON -0.10662 Leonberger
LHAS 0.121432 Catahoula Leopard Dog
MALT 0.133881 Maltese
MANT 0.03463 Manchester Terrier (Standard)
MAST 0.0893 English Mastiff
MBLT 0.242318 Bull Terrier Miniature
MNTY 0.270766 Manchester Terrier (Toy)
MPIN 0.115542 Miniature Pinscher
MPOO 0.090096 Poodle small Miniature
MSNZ 0.150345 Schnauzer Miniature
NELK 0.010105 Norwegian Elkhound
NEWF 0.085381 Newfoundland Lanseer
NORF -0.0333 Norfolk Terrier
NOWT 0.180618 Norwich Terrier
NSDT 0.10002 Nova Scotia Duck Tolling Retriever
OES 0.11044 Old English Sheepdog
PAPI 0.118812 Papillon
PBGV 0.056894 Basset Griffon Vendeen Petit
PEKE 0.065984 Pekingese
PEMB 0.046101 Pembroke Welsh Corgi
PHAR 0.258646 Pharaoh Hound
PNTR 0.127687 English Pointer
POM 0.190252 Pomeranian
PRES 0.170424 Presa Canario
PTWD 0.157522 Portuguese Water Dog
PUG 0.259613 Pug Dog
PULI -0.11027 Puli
RHOD 0.20009 Rhodesian Ridgeback
ROTT 0.087514 Rottweiler
SALU 0.148652 Saluki
SAMO 0.097442 Samoyed
SCHP 0.063912 Schipperke
SCOT 0.056301 Scottish Terrier
SCWT 0.303182 Soft Coated Wheaten Terrier
SD-DIST 0.065265 Sled Dog – Distance
SD-SPR -0.20197 Sled Dog – Sprinter
SHAR 0.248821 Chinese Shar Pei
SHIB 0.158441 Japanese Shiba Inu
SHIH 0.155844 Shih Tzu
SILK 0.075987 Silky Terrier
SPIN 0.108583 Italian Spinone
SPOO 0.140248 Poodle Standard
SSHP 0.140501 Shetland Sheepdog
SSNZ -0.03506 Schnauzer Standard
STAF 0.024789 Staffordshire Bull Terrier
STBD 0.104228 Saint Bernard
SVAL 0.083595 Swedish Vallhund
TIBS 0.066005 Tibetan Spaniel
TIBT 0.215652 Tibetan Terrier
TPOO 0.002949 Poodle small Toy
TURV 0.024285 Belgian Tervuren
VIZS 0.040201 Hungarian Viszla
WEIM 0.064422 Weimaraner
WELT 0.117925 Welsh Terrier
WHIP 0.113562 Whippet
WHPG 0.072099 Wirehaired Pointing Griffon
WHWT 0.096207 West Highland White Terrier
WSSP 0.236838 Welsh Springer Spaniel
YORK 0.159496 Yorkshire Terrier
You have to be careful about Turnbull’s Blue. Blue may have been an old bearded farm collie crossed to a Border Collie (which was popular) or simply a “bearded” type of border collie. Also Blue is a “ROM” with the ISDS and also …. registered with the Kennel Club as a Bearded Collie and is in their registry so he is dual listed!!!
This article says nothing about outcrossing to other breeds being a good thing. How is it productive unless it’s for a good reason, like let’s say creating a line of clears for an extremely common problem, like urine crystals in Dals. Otherwise its only effect is creating a smaller COI. Which is only a number.
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Um, so what? This article is about what it’s about. It’s not the only one on this site. There are several hundred more, have fun reading some of them. The goal here is to assess the degree to which Border Collie breeding mirrors Alaskan Sled Dog breeding given published analysis of others that indicates a fondness for that analogy. Whatever implications of the outcrossing are on health or performance are secondary and implied by the argument I’m responding to.
Well, “productive” depends on what your goals are, short and long term. If you are married to the idea of blood purity and closed registries for their own sake, then outcrossing would NEVER be “productive” to your highest goals, even if it provided myriad benefits for other goals which are perhaps more important or valuable.
As for outcrossing in general, introducing a healthy allele is of course a benefit that is productive if health is your goal. But that is not the only benefit. Consider eating and drinking in the absence of hunger and starvation and disease. Are those activities “productive”? Should you have to wait until your kidneys shut down before you drink water every time? Should malnutrition be the only indication to eat food? Is it wise to consume vitamins routinely or only if you’ve been diagnosed with a deficiency that’s leading to a pathology?
Heterozygosity itself has advantages all by itself. Not only does it often prevent expression of deleterious mutations that exist in the breed and which arise often from routine mutations each generation, it allows breeders a greater palette of traits to select from in forming their breeding programs.
Consider this, as breeders we could scour the entire population of dogs to find the perfect male and the perfect female. We could then declare that those two dogs are the ideal and then limit the breed to just those two dogs. And of course their offspring. Why do we not do this? Well, because no one judge’s opinion is supreme on what makes their ideal dog. So diversity is inherent in the system due to the diversity of values and traits and opinions. The only way to satisfy those competing values and maximize the utility we gain as a group from them is to allow for as much diversity as is wanted.
Breeds that are not defined by expressly axing diversity are very often very diverse across traits and yet people can still form mental models of them.
So no, it’s not about COI and no heterosis is not simply a number. That’s a very poor reflection of your understanding of both genetics, evolution, population structure, heterozygosity, inbreeding, etc. It might be “only a number” to you. But that’s as banal and ignorant as saying π is “only a number” as your sole assessment of the entire field of math.
COI is a part of how we talk about breeding structure, but it’s not the only thing and it’s certainly not as central or as limited as you seem to be implying.
Reintroducing lost heterozygosity or reducing the frequency of undesirable genes (traits) is not a ‘good reason,’ Christopher.
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I think this article gets stuck in the weeds a bit and misses the most important point on the Border Collie/Alaskan Husky comparison (I’m an Alaskan and know or have known multiple Quest and Iditarod mushers and have a fair idea about their breeding programs).
The point, in my opinion, is breeding for performance, behavior and health as opposed to conformation and trotting around a show ring as the ultimate measure of the dog’s value.
Hopefully, for the dogs’ sake, Border Collie breeders stay focused on the former!