Inbreeding leads to:

Increased homozygosity.

Increased expression of recessive traits.

Decreased expression of dominant traits.

Increased heterozygosity.

An allele:

Is one of possibly many variations of a gene.

Can be recessive.

Can be dominant.

Is a synonym of “gene.”

A gene:

Is a unit of heredity that is transferred from parent to offspring.

Controls the transmission and expression of one or more traits.

Can code for multiple and otherwise unrelated phenotypic effects.

Codes only for a single discernible phenotypic effect.

Can be dominant or recessive.

If an individual has inherited the same allele for a gene from both parents, they are said to be:

Homozygous.

Hemizygous, Nullizygous, Allozygous, Heterozygous.

Phenotype refers to:

The set of observable characteristics of an individual.

The morphology, behavior, and development of an individual.

The genetic makeup of an individual.

Both expressed and unexpressed genetic variation.

Which of the following trait types can “breed true?” To “breed true” means that two organisms with a particular heritable phenotype produce only offspring with that same phenotype.

Recessive traits. (e.g. brown coloring, rough coats, tri-color, etc.)

Dominant traits. (e.g. black coloring, smooth coats, etc.)

Heterozygous Semi-Dominant traits. (e.g. Merle, Bobtail, Chinese Crested Hairless, German Shepherd Panda, etc.)

Heterozygous Co-Dominant traits. (e.g. AB blood type, red-white Roan in horses, etc.)

What mode of inheritance do the following traits have in common: Merle, Bobtail, Harlequin, Panda (German Shepherds), Hairlessness (Xolo, Chinese Crested, etc.) ?

They are Lethal.  Lethal genes are capable of causing death.

They are all semi-dominant. When the heterozygote has a different, intermediate phenotype compared to the homozygous dominant or homozygous recessive individuals, it is said to be “semi-dominant.”

They are all recessive.

They are all dominant.

They are sex-linked.

They are all co-dominant.

Inspired by the Genetics Survey on Gene Expression and an inquiry from the CanineGenetics-L yahoo group, I figure it might be fun to post an occasional genetics quiz to reinforce some of the concepts covered on this blog and provoke discussion of the sorts of genetics knowledge that is most helpful to dog enthusiasts.

The results are not tracked, it’s completely anonymous, and the correct answers will be revealed tomorrow. I tried several automated quiz plugins but they are a pain, so we’ll just go old school.  If you run across an answer that you want to discuss, do leave a comment.

February is the month of sweethearts, so there’s no better time to bone up on what happens when love is in the air and dogs are swapping genes. For the inaugural monthly quiz, here are some questions long time readers of the blog should be able to answer.

Multiple Correct Answers are accepted and often required.

Inbreeding leads to:

Increased homozygosity.

Increased expression of recessive traits.

Decreased expression of dominant traits.

Increased heterozygosity.

An allele:

Is one of possibly many variations of a gene.

Can be recessive.

Can be dominant.

Is a synonym of “gene.”

A gene:

Is a unit of heredity that is transferred from parent to offspring.

Controls the transmission and expression of one or more traits.

Can code for multiple and otherwise unrelated phenotypic effects.

Codes only for a single discernible phenotypic effect.

Can be dominant or recessive.

If an individual has inherited the same allele for a gene from both parents, they are said to be:

Heterozygous.

Homozygous.

Hemizygous.

Nullizygous.

Allozygous.

Heterozygous.

Phenotype refers to:

The set of observable characteristics of an individual.

The morphology, behavior, and development of an individual.

The genetic makeup of an individual.

Both expressed and unexpressed genetic variation.

Which of the following trait types can “breed true?” To “breed true” means that two organisms with a particular heritable phenotype produce only offspring with that same phenotype.

Recessive traits. (e.g. brown coloring, rough coats, tri-color, etc.)

Dominant traits. (e.g. black coloring, smooth coats, etc.)

Heterozygous Semi-Dominant traits. (e.g. Merle, Bobtail, Chinese Crested Hairless, German Shepherd Panda, etc.)

Heterozygous Co-Dominant traits. (e.g. AB blood type, red-white Roan in horses, etc.)

What mode of inheritance do the following traits have in common: Merle, Bobtail, Harlequin, Panda (German Shepherds), Hairlessness (Xolo, Chinese Crested, etc.) ?

They are Lethal.  Lethal genes are capable of causing death.

They are all semi-dominant. When the heterozygote has a different, intermediate phenotype compared to the homozygous dominant or homozygous recessive individuals, it is said to be “semi-dominant.”

They are all recessive.

They are all dominant.

They are sex-linked.

They are all co-dominant.

Answers Here

Champis, the Swedish herding rabbit.

Move over Barbie Collies, there’s a new herder in town that is adorably small, has perfect ear set and has a decidedly fluffy butt.  And he’s not a show Border Collie from Australia, he’s a rabbit from Sweden.

Nils-Eric and Greta employ their granddaughter Hanna’s rabbit named “Champis” on their farm in Käl, Sweden to help herd their sheep. It appears that Champis learned to herd sheep–complete with heading, heeling, eye, and clapping behaviors–from their Border Collie, but this video shows that Champis gets along just fine all by himself and the sheep respect his authority.

"With a rabbit like Champis, who needs a dog?"

Frankly, I’m amazed that the rabbit seems to know what the shepherd wants, just like the Border Collies do; for example, he runs to cut the sheep off from re-entering the barn and he even makes a valiant effort to bring back a sheep that has gone stray, and he’s perfectly willing to stare down a bull headed woolly.

..

Read the continuing adventures of Champis the herding rabbit (named after a Swedish soft drink) and his Border Collie friends “Gimmie” and “Fame” on the Gårdsbackens blogg.

The retina of a Briard with an inherited blindness disease that was later cured with gene therapy.

Back in 2001, scientists first treated inherited blindness in dogs using gene therapy. In that instance it was Congenital Stationary Night Blindness which manifests in Briards and is analogous to Leber congenital amaurosis in humans, both defects in the RPE65 gene.

Dogs blinded by an inherited retinal degenerative disease had their vision restored after treatment with genes from healthy dogs, marking the first successful gene therapy for blindness in a large animal. The treatment offers hope for humans with a similar condition.
…
“We have shown that gene therapy can restore vision in dogs with one of the most clinically severe retinal degenerations,” says Acland, a research veterinarian at Cornell’s James A. Baker Institute for Animal Health.
…
The RPE cell layer in the eyes of humans, dogs and other mammals supports the retina by providing nourishment and removing waste products while supplying vitamin A to the photoreceptors. Puppies and human infants with defective RPE65 genes produce a mutant form of the RPE65 protein, resulting in early vision loss, degeneration of the retinas and near-total blindness later in life.

The canine form of this retinal degenerative disease has been found only in the briard dog breed. In the gene therapy experiments, researchers used RPE65 genes that were cloned from dogs without the disease, together with a viral vector (recombinant adeno-associated virus, or AAV) to carry the normal dogs’ DNA. They injected the combination into the subretinal space of the eyes of 3-month-old briard-beagle mix dogs that were known to have the defective RPE65 gene and had been blind since birth. Within six weeks, the treated eyes were producing the correct form of RPE65 protein. By three months, a series of tests (electroretinography, pupillometry and obstacle-avoidance tests in a dimly lit room) demonstrated that vision was restored to the treated eyes.

In 2010, the same team successfully returned function to the cones of previously blind dogs affected by a form of blindness called Achromatopsia.

Now, scientists have again used gene therapy to prevent and reverse another form of blindness in dogs and humans called X-linked Retinitis Pigmentosa associated with a mutation in the RPGR gene.

The disease in humans and dogs is caused by defects in the RPGR gene and results in early, severe and progressive vision loss. It is one of the most common inherited forms of retinal degeneration in man.

“Every single abnormal feature that defines the disease in the dogs was corrected following treatment,” said lead author William Beltran, assistant professor of ophthalmology at Penn’s School of Veterinary Medicine.

“We were thrilled,” said senior author Gustavo Aguirre, professor of medical genetics and ophthalmology at Penn Vet. “The treated cells were completely normal, and this effect resulted from introducing the normal version of the human gene into the diseased photoreceptor cells.”

This is exciting for dog owners because there are numerous breeds that have epidemic levels of eye disease and the proliferation of these advancements could lead to routine treatments that would treat or cure the disease in individuals.

There’s still a barrier to a permanent cure for breeds that would extend to the offspring of treated individuals.  Treated dogs only receive the working DNA in the somatic cells at the location of the viral vector.  Somatic cells are all the cells in the body besides those directly involved in producing offspring.  If there’s a mutation in a somatic “body” cell, it will not be passed on to future generations.

The cells that produce the sperm and egg are called germline cells and they are not necessarily amenable to the same gene therapy techniques that work on individual somatic cells.  You can effectively cure the blindness by providing a working copy of the genes within the eye, but other cells in the body will not have the new gene and the germline cells will not produce sperm or eggs that benefit from the disease-free allele either.

This effect is called the Weismann barrier, which is the theory that genetic information moves only in one direction, from germline cells to somatic cells.  This is why when you get an x-ray the tech is more concerned with lead shielding your private parts than the rest of your body because germline mutations have serious consequences for your offspring and germline cells are theoretically immortal (they can replicate for the entire life of the organism) whereas somatic cells only divide 30-50 times.

If the Weismann barrier can be broken, it’s theoretically possible for the treatment of eye cells to result in genetic change in somatic cells and thus in the offspring.

There’s also the potential to apply gene therapy techniques to germline cells (such as the sperm or egg) directly and this would enable genetic engineering that would last over generations.  Scientists have already created lasting changes in organisms that do not differentiate somatic-germline cells (like plants: thus we have GMOs), but I’m not aware of any ongoing progress on germline gene therapy and there are currently institutional barriers against work in that area given the potential ethical implications of genetic engineering.  Scientists have successfully inserted synthetic chromosomes into mice that were heritable but the current focus in treatment of human offspring disease is to perform IVF and genetically profile the blastocysts before implantation.

So this is good news for dogs, and possibly the beginning of an amazing new world of animal husbandry when genetic engineering will allow for some truly incredible health advancements and possibly some radical experimentation.

3 minutes a day in a centrifuge was part of the failed biosensor "super dog" program.

Dog culture is lazy and unoriginal, and profoundly stagnant. The desire for easy answers, simplistic how-tos and formulas for success is rampant. Just do this one simple thing! 

Breeders eschew complexity, uncertainty, and experimentation. They fear change and embrace unproven tradition on face value.  We do it this way because we’ve always done it this way.

Reason gives way to mimicry, and that is the true mark of conformation: not in the dogs keeping to a written standard but in breeders kowtowing to the unwritten rites and rituals to fit in.

One cherished ritual that can be found proudly advertised on numerous breeder websites (usually after the “Our Boys” and “Our Girls” links) as a sign of their reputable status and deep commitment to superior dogs is the adoption of the “Bio-Sensor” program as the one true path™ to dog raising.

In only 15-25 seconds a day for only 14 days in a dog’s life you will realize “life long lasting effects:” “improve performance,” “respond maximally,” “attain sexual maturity sooner,” “resist cancer and infectious disease,” “withstand terminal starvation,” achieve “psychological superiority,” “stronger heart beats,” “stronger adrenal glands,” and “improved cardio vascular performance!”

That’s less than 6 total minutes of work to make a super dog!  AMAZING!

The brains over at the sci-ence blog have come up with a handy chart to recognize quackery, the relevant parts of which I’ve reproduced here.  Their instructions:

“If you come upon a treatment or product that seems too good to be true, consult this handy guide to finding pseudoscience, scams, and quack medicine. Remember, it only takes one match to be considered suspect! Be safe, and be skeptical!”

This widely tauted “Bio Sensor” a.k.a “Super Dog” a.k.a “Early Neurological Stimulation” program has many warning signs of quack science.

Let’s start with who is peddling this pseudoscience:  Dr. Carmen a.k.a Carmelo Battaglia, Board of Directors of the American Kennel Club.

Dr. Carmen L. Battaglia, of Roswell, Georgia, owns and breeds German Shepherd Dogs and is Delegate and Past President of the German Shepherd Dog Club of America. Carmen has chaired the Committee for the Future and Business/Planning Committee and as a former AKC Director, served as Board liaison for the Health, Parent Club, HEC and By-Laws Delegates committees. He has published articles on breeding and legislation as well as several award-winning books. He also serves as an AKC expert witness in dog legislation cases and has written county dog legislation which resulted in the model that is used in several states. Carmen possesses a Doctorate from Florida State University and has been Assistant Dean at Emory University and Regional Administrator at the US Department of Education. He is also the President of Atlanta Student Aid (financial aid consulting Firm) as well as the past president/owner of three post secondary schools which were located in two states.

You’ll remember Dr. Battaglia from his resurrection of Lloyd Brackett and his infamous “Brackett’s Formula.”  Dr. Battaglia gives lectures on cruise ships to up-and-coming brown-nosers in the AKC hierarchy who want to buy the secret knowledge and pay the right gate keepers to fast track show success.  He’s the closest thing the AKC community has to a celebrity doctor.

But don’t get too comfortable with the idea that he’s a medical doctor, he isn’t.  He’s a Ph.D. doctor, which he readily advertises at the end of his publications:

Carmen L Battaglia holds a Ph.D. and Masters Degree from Florida State University. As an AKC judge, researcher and writer, he has been a leader in promotion of breeding better dogs and has written many articles and several books.

Dr. Battaglia is also a popular TV and radio talk show speaker. His seminars on breeding dogs, selecting sires and choosing puppies have been well received by the breed clubs all over the country. Those interested in learning more about his seminars should contact him directly.

Visit his website at http://www.breedingbetterdogs.com

What he doesn’t advertise anywhere that I’ve found despite an extensive search is what subjects his degrees are in.  So I contacted the Curriculum Publications Coordinator at Florida State University and found out the unpublished truth:  B.A. Psychology 1958, M.S. Social Welfare 1960, PhD Joint Doctoral Program in Criminology Corrections and Sociology 1968.

So by way of education, Dr. Battaglia is more equipped to run a prison than a breeding program.  His dissertation was titled “Deviant behavior of parolees and the decision-making process of parole supervisors.”

The next indicator that “Bio-Sensor” is quack science is because it’s being SOLD as a how-to guide to success without having been vetted in any way by scientists in peer reviewed publications resulting from studies done according to the scientific method.

Dr. Battaglia sells his program along with breeding and puppy selection advice as part of his self-help for dog breeding commercial venture.  You can buy books, videos, DVDs, and subscribe to his newsletter and attend his lectures.

In accordance with yet another quackery red flag, Dr. Battaglia is pitching program that offers medical benefits but he (nor anyone else) has no peer-reviewed journal articles on his protocol. And it’s not for lack of trying.  If you visit his website you will find a link to request his so far unpublished journal article.  When you do so, you will be e-mailed a copy of an extended version of his Bio-Sensor article spruced up to look like an actual experiment with “Methods and Materials” and everything.

It appears from the file that Dr. Battaglia has attempted to get this article published since at least 2007, but he will warn you that the article is still under intense review and thus you can not share it.

This wouldn’t be the first time that the “Bio Sensor” program has been used to sell a self-help program, however, as Dr. Battaglia collaborated with Stanley Coren–king of marketing shoddy dog science to pet owners in book form–who included the information in his book Why Does My Dog Act That Way?

Battaglia and Coren’s considerable influence on the dog fancy combined with Battaglia offering the super simplified how-to instructions for achiving super dog success for free on his website as a teaser for his suite of videos, books, and lectures the “Early Neurological Stimulation” program has saturated the hobby pet breeder culture.  Breeder testimonials and reprints of the method are everywhere.

Diligently following Dr. Battaglia’s advice, there are breeders out there inbreeding their lines and producing singleton puppies who none-the-less credit Brackett’s Formula and the amazing Bio-Sensor program for giving them a super puppy abounding with exceptional qualities.

Apparently I’m part of a silent minority who have even questioned this program’s merits and the academic bona fides of the man who peddles it from coast to coast, as I’ve found no online criticism of the methods and not even one other soul who questioned what Dr. Battaglia’s field of study was until I sent out feelers over a year ago.

But it’s clear to me that Bio-Sensor is being marketed just like quack science is marketed, by people who have a vested financial interest in selling easy answers and quick fixes to gullible pet breeders who spend fortunes trying to rectify their ignorance with short cuts and feel-good nonsense.

In a quadrumvirate of quackery, Battaglia also uses his “Breeding Better Dogs” website to sell Japanese (think Eastern medicine) magic magnetic shoe inserts and magic magnetic dog beds that are “combined with magnetic technology, another ancient principle” and “enhance the body’s energy flow to allow healing and proper metabolism.”  This is a man perfectly willing to market quack science as a miracle product for profit.

So far I’ve shown that Bio-Sensor looks like a duck, and in a future post I’ll show you how it quacks like a duck as well.

The breed is believed to be the creation of one man, Heinrich Essig, who was a dog dealer and traveling salesman; his claimed formula was Landseer Newfoundland x Saint Bernard, followed by more Saint Bernards and a Pyrenean Mountain Dog with the goal being a large white dog that was fashionable at the time.  After Essig’s death in 1889 his nephew had the inspiration to promote the breed as a lion-like mascot for the town of Leonberg and established the final conformation of the dogs as having a lion-like rough coat with reddish-brown coloration, a black mask and black sable accents.  Like most breeds, the origin mythology is poorly documented and stud books were not kept in earnest on the breed for several decades after its establishment.

As a German breed, the Leonberger was severely affected by both World Wars: only 5 known breeding dogs survived the first war and only 8 pedigreed dogs emerged from the second.  Two severe bottlenecks like this are not conducive to preserve genetic diversity within a closed population.

That is why I was surprised to find that the 5 Leonbergers which were DNA tested as part of a study analyzing the genetic composition of Alaskan Sled Dogs showed that they had excess heterozyosity compared to Hardy-Weinberg equilibrium. Of 141 breeds tested over 96 genetic marker sites, there were only 9 purebred breeds found to be surplus heterozygous and most of those are only marginally so.

Note, this is not a measure of gross diversity in the breed, it’s a measure of which way the breed is being pushed genetically, either toward more conformity and homozygosity or toward more diversity and heterozygosity.

Genetic rescue requires a push toward heterozygosity before reaching a new equilibrium with the new alleles.

Inbred or Outcrossed? Bars that extend to the left indicate excess heterozygosity compared to the Hardy-Weinberg equilibrium (red line), suggesting active selection for allele diversity (i.e. outcrossing). The Leonberger is one of the most heterozygous breeds tested.

Here is the actual data represented in the chart.  The purposely out-crossed hybrid sled dogs were found to be 20% more diverse than equilibrium and the Leonberger was second only to the Puli with over 10% excess heterozyosity.

I’ve only found one documented outcross on the Leonberger books: In 1954 a Leonberger stud Arko von Leonberg with a COI of 20% was bred to a Newfoundland dam Grisette von Bruckberg and one female from that litter, Alma von Rossbach, would cement herself into the Leonberger gene pool.

More recently European kennel clubs have registered Leonbergers with full breeding rights certificates “titre initial” (as opposed to a 3 generation provisional appendix registration “registre initial”) which are given after a number of criteria are met; typically an evaluation against the breed standard or significant show success, health testing, and perhaps even temperament evaluations.  These dogs are believed to be pure-blooded Leonbergers instead of hybrid dogs which would normally take 3 generations to be admitted into the gene pool with full breeding rights and purebred status in registries that allow new blood.  Sometimes they come with full pedigree and while the kennel club would void the known pedigree and perhaps calculate the COI of offspring as 0%, it’s not clear that either the registre initial or titre initial schemes are a source of new Leonberger blood.

Another possible source of Leonberger diversity lies in the history of Germany and the East after the wars.  The Leonberger isn’t just a dog of the West and so it existed on both sides of the Iron Curtain.  It is unknown and undocumented what sorts of possible crosses came into the breed during the Cold War before the reunification of Germany and the resumption of normal trade across Europe.

The last and least verifiable source of unexpected diversity would be intentional and unintentional pedigree fraud or error.  It’s possible that some breeders when faced with a line that was not producing what they wanted or suffering from inbred disease or infertility decided to outcross, or an accidental litter proved sufficiently virtuous to register and the breeder either could not or did not choose to disclose this.

There’s also the possibility that I simply haven’t accounted for some factors which would have preserved heterozygosity in Leonbergers and that the bottlenecks did not have the expected effects on the gene pool.  Certainly more questions than answers, but what an intriguing mystery it is.

Hot Nun knows a species when she sees one. Obey the Sister.

After the smashing success of my Multiple Orcasms post which still brings considerable daily traffic to the blog from furries looking for orca and vore themed pornography, I couldn’t resist tiptoeing around cheeky references to bestiality once again; but this time the human interest in animal sex is strictly like-on-like and the link to pornography is in the tricky means of defining concepts that are both familiar and yet abstract.

Trying to define what is necessary and sufficient to designate a “species” is rather like the problem the US Supreme Court ran into when trying to define pornography.  In the decision for Jacobellis v. Ohio, Justice Stewart coined a now famous phrase when trying to draw a line between protected speech and unprotected obscenity:

In saying this, I imply no criticism of the Court, which in those cases was faced with the task of trying to define what may be indefinable… I shall not today attempt further to define the kinds of material I understand to be embraced within [hardcore pornography]; and perhaps I could never succeed in intelligibly doing so. But I know it when I see it, and the motion picture involved in this case is not that.

Many definitions, while concrete, are rather relative and not absolute. Darwin failed to define species in his Origin of Species:

Nor shall I here discuss the various definitions which have been given of the term species. No one definition has as yet satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species.

He goes on to say just how blurred that line can be:

I was much struck how entirely vague and arbitrary is the distinction between species and varieties.

The “species debate” is a problem that existed before Darwin and which continues today.  The current tone of the debate has been most significantly influenced by evolutionary biologist Ernst Mayr whose definition of a species graces most modern textbooks:

species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups

I had a laugh when I read Ernst Mayr’s essay “What is a Species, and What is Not?” published in Philosophy of Science in 1996:

The term ‘species’ refers to a concrete phenomenon of nature and this fact severely constrains the number and kinds of possible definitions. The word ‘species’ is, like the words ‘planet’ or ‘moon,’ a technical term for a concrete phenomenon. One cannot propose a new definition of a planet as “a satellite of a sun that has its own satellite,” because this would exclude Venus, and some other planets without moons. A definition of any class of objects must be applicable to any member of this class and exclude reference to attributes not characteristic of this class.

It doesn’t help Mayr’s point that the International Astronomical Union had no formal definition for ‘planet’ at the time of his essay or for a complete decade after it. When they did finally vote on one in 2006, the former planet Pluto got downgraded to a dwarf planet leaving every science textbook published in the preceding 80 years obsolete and violating Mayr’s apparent rule that one can not propose a new definition that would exclude an existing member of a class. Pluto was declared a planet upon its discovery in 1930 and now it’s not even the largest “plutiod”–that honor falls to Eris, which was discovered in 2005 and found to be larger than Pluto which prompted Astronomers to actually look at the definition of what makes something a planet versus something else.

If you take a moment to look around the blogosphere this week, you’ll realize that the uncertainty of this issue is present just beneath the surface of numerous topics of conversation:

Retrieverman asks if the Island Fox is a valid species which must be understood within the context of the greater Canid complex several species of which would violate the basic Mayr definition of a species, but his post on the Polar Bear is also framed by the species debate as genetic analysis shows that Polar Bears can be considered a variety of Brown Bear and the two can and do form fertile hybrids.

The Dog Zombie looks at Canid DNA to question the recipe of different flavors of “Canid Soup.”

Jess at Desert Wind Hounds picks up on the food metaphor at Dog Zombie and asks what the recipe is for a “purebred” and how one goes about creating one.  This is ultimately just a more zoomed in analysis of the species debate: how much distance in time, space, genetics and niche constitutes a different breed, a different type, a different landrace, or a different species?

Stephen Bodio asks the same question with a simple image comparison of two dogs.

Razib Khan shows that the species debate is applicable to humans, and the notion that Neanderthals or Denisovians were somehow not human falls when you realize that their genes are still in us (Border Wars is written by a 2.7% certified Neanderthal):

In my post below I argue that it’s most useful to reconceptualize “human” as an ecological niche, rather than a descent group. All the confusion as to whether Neandertals, or any other group of divergent hominins, were, or weren’t, “humans like us,” exists in the context of the idea that “humans like us” are a very specific and sui generis  cladewith special traits. I think “we” need to get a little off our high horse here.

You’ll notice that the notion of “niche” becomes more and more important as we realize just how blurry the lines between interfertile species really is.  Niche is what separates Polar Bears from Brown Bears and it’s also what separates Dogs from Wolves:

The co-evolution between social canids and primates is I think not a random chance event. To some extent I think “man’s best friend” was a necessary outcome of evolutionary forces. Barring the total extermination of one lineage or the other, some sort of cooperative relationship is I suspect something that will naturally reoccur. Dogs are not simply a specific derived lineage of wolves, they’re an ecological niche created by the existence of hominins with social complexity.

Dave at Prick-Eared has a post which documents another canid rapidly invading the very niche that once brought man and wolf together to co-evolve.

There are no hard and fast answers here, no absolute definitions, no minimum standards or list of traits that are both necessary and sufficient to differentiate one “thing” from another “thing” in a meaningful way.  This is the place where the objectivity of science meets the subjectivity of philosophy and those questions like “what is a dog” start to look a lot like “what is an ideal Afghan Hound.”

There are some questions that are worth answering “I don’t know and I probably never will, but that won’t stop the investigation.”

The wolves of Isle Royale Michigan

Sweeping comparisons between wild animals and domesticated pets are dangerous given the unique and often mutually exclusive conditions in which those two groups often find themselves.  Wild animals must hunt for their food, compete for resources, suffer predation, lack medical intervention, self-select their mates, and exist in numbers based upon a complex interaction between their own merits and the conditions in their environment. Pet dogs are fed from a bowl daily, compete only for our affection and attention, are largely free of predation, have ready access to modern veterinary techniques and treatments, have their mates chosen for them–sometimes from dogs long dead or on other continents, and exist in numbers based upon human concerns and rarely on their own merits or the environmental carrying capacity.  Their selection is very much unnatural. But there are very few scientific studies of domesticated pets versus numerous investigations into wild populations, so dog lovers would be remiss in not learning lessons from our pets’ wild cousins.  One particularly interesting ongoing scientific inquiry is the study of the wolf and moose populations on Isle Royale Michigan.

Wolves colonized Isle Royale, a wilderness island in Lake Superior, North America, in 1949 or 1950. The population is isolated from mainland wolves by a channel of frigid water, 24 km wide. In many, but not all years, this channel freezes for several days or weeks. Although an occasional ice bridge makes immigration possible, the analysis of mitochondrial DNA and the Y chromosome suggests that the population was originally founded by only one female and two males.

This makes Isle Royale analogous to a small dog breed based on a few founders or even a single kennel that rarely brings in any new blood.  Nova Scotia Duck Tolling Retrievers, Leonbergers, Cavalier King Charles Spaniels, Vallhunds, and many other breeds were founded by only a few sires and dams and have had very little influx of blood since and there are quite a few old time breeders who would only occasionally bring in a sire from outside their own moderately sized kennels. As with the above breeds, the mostly-closed gene pool on Isle Royale became steadily inbred over time.

By the late 1990s, the population’s estimated inbreeding coefficient had risen to 81%. Fifty-eight per cent of Isle Royale wolves showed congenital bone deformities compared with only 1 per cent in two outbred wolf populations. Some of these deformities could reduce individual fitness, particularly components of fitness associated with predation and reproduction.

So the first lesson to be learned from the Isle Royale Wolves is that isolation leads to inbreeding and inbreeding is detrimental.  In dogs, closed registries and kennel blindness are a form of isolation and in both cases we see rising inbreeding followed by increased expression of otherwise rare diseases.

Some of the most studied wolf packs in the world are in serious jeopardy. Researchers report that the occurrence of debilitating bone deformities in wolves marooned on Isle Royale, an isolated island in Lake Superior north of Michigan, has risen sharply over the past five decades due to inbreeding.

Diseased wolf vertebra due to inbreeding of Isle Royale wolves.

A genetic defect now common in the Isle’s wolves causes bones in the spine, the vertebrae, to grow gnarled and crooked. Also found in domestic dogs – close wolf relatives – the bone malformations  can pinch nerves in the spinal cord, causing pain that makes it tough to walk and can lead to paralysis of the back legs and tail in severe cases, according to research published in February’s issue of Biological Conservation. Back in the 1960s, about a quarter of Isle Royale’s wolves appeared to have the anatomical abnormality, but now the percentage of afflicted wolves has risen to nearly 60 percent of the population. “In normal, healthy wolf populations without inbreeding, you are only supposed to see this kind of defect in about one out of a hundred animals,” says paper coauthor John Vucetich, an assistant professor of wildlife biology at Michigan Technological University (MTU) in Houghton. The deformity, discovered during autopsies of recovered, dead wolves, has grown so rampant, Vucetich says, “we haven’t found a normal wolf in the past decade.”

- Scientific American

The second lesson to learn from the Isle Royale Wolves is that hybrid vigor is real and powerful and that inbreeding depression is real and powerful.

We used molecular techniques to document the consequences of a male wolf (Canis lupus) that immigrated, on its own, across Lake Superior ice to the small, inbred wolf population in Isle Royale National Park. The immigrant’s fitness so exceeded that of native wolves that within 2.5 generations, he was related to every individual in the population and his ancestry constituted 56 per cent of the population, resulting in a selective sweep of the total genome. In other words, all the male ancestry (50% of the total ancestry) descended from this immigrant, plus 6 per cent owing to the success of some of his inbred offspring. The immigration event occurred in an environment where space was limiting (i.e. packs occupied all available territories) and during a time when environmental conditions had deteriorated (i.e. wolves’ prey declined).

This event is an excellent example of what one outcross might do for a small breed and especially a single kennel. The immigrant wolf’s offspring were true hybrids, a mix of the formerly isolated pool and fresh blood; versus having an outside pack supplant the locals in toto, as one might expect in a contiguous geography that was not isolated like the Isle.

We have here an analogous situation that fits existing human patterns of behavior regarding dog breeds and strains that have virtual barriers instead of physical ones.

The astounding aspect of this immigrant on Isle Royale is just how potent his genetics were to effect change on the island.  None of the existing males could even compete with him and he became the sole sire.  His initial success is likely caused by the inbreeding depression in the inbred Isle wolves.

The high fitness of this immigrant wolf was also associated with distinctive behaviour and physical appearance. First, he was physically larger than most Isle Royale wolves. As alpha male of the Middle Pack, his high fitness was also reflected by his dominance over other ISRO packs. Specifically, he exhibited strong territorial behaviour that completely displaced West Pack, driving that pack to extinction by 1999.

This immigrant wasn’t necessarily a super-wolf, he was in all probability a young male that was driven out of his birth pack’s territory for being non-competitive with that pack’s alpha male. His success on Isle Royale shows just how compromised the inbred population had become.

The inbred Wolves of Isle Royale, genetically refreshed by the genes of a new sire.

It’s an open question if and how much the influx of new genes has changed the bone deformities which had come to define Isle wolves.  Surprisingly, before the results of the bone study were published, Isle Royale was used as an example of a wild population that was thriving and unharmed by inbreeding and isolation.  This is why I’m cautious of anyone who argues from ignorance regarding their ability to inbreed and avoid disease.  This is the third lesson: don’t assume that inbreeding can exist in high levels without detriment and don’t cite wild populations if no one has ever done a detailed health study to document the true health of the population.

The new results offer the first evidence of the wolves’ closed population leading to a decline in natural fitness. This is important, Vucetich says, because for years some policy makers and conservationists have pointed to the apparent health of the Isle Royale wolf packs as an indication that small animal populations can maintain proper genetic diversity. “Isle Royale is not this robust place that some people thought it was,” says Vucetich.

Now, not all the lessons are positive ones.  Given the isolation of the island, the complete genetic sweep of the immigrant male, and the small population size the rates of inbreeding swiftly ticked back up.  The smaller the population size and the greater degree of inbreeding done following new blood, the shorter time you’re going to reap the benefits of that new genetic material.  Outcrossing can forgive a myriad of sins, but it needs to be used in measure to the problem.  On Isle Royale, this new wolf didn’t simply add to the sires on the island, he supplanted them and then bred with his children, creating more inbreeding instead of extending the benefits of outcrossing.

This isn’t an advisable strategy for refreshing a dog breed, essentially going from one popular sire to another.  Diversity in breeding males should be maintained over each generation and proceeding to breed father to daughter–and similar–after a single outcross will swiftly return the gene pool to inbred status. The other issue that researchers have is that to complete a “genetic rescue” one must be able to document the benefit on a population level, an this is often done with the most crude methods, namely demography (head counting).  An increase in population size was not observed here, although there was a precipitous drop in the wolves’ main food source, Moose, during this time and the wolf numbers did not suffer either.

Genetic rescue, in which the introduction of one or more unrelated individuals into an inbred population results in the reduction of detrimental genetic effects and an increase in one or more vital rates, is a potentially important management tool for mitigating adverse effects of inbreeding. … The immigration event occurred in an environment where space was limiting (i.e. packs occupied all available territories) and during a time when environmental conditions had deteriorated (i.e. wolves’ prey declined). These conditions probably explain why the immigration event did not obviously improve the population’s demography (e.g. increased population numbers or growth rate). Our results show that the beneficial effects of gene flow may be substantial and quickly manifest, short-lived under some circumstances, and how the demographic benefits of genetic rescue might be masked by environmental conditions.

This is not a problem with outcrossing at all, rather it’s a problem of the limitations of the crude science of demography. Head counting might speak to quantity, but it is wholly lacking in useful information about quality.

Demography is likewise a poor tool to employ when analyzing domesticated dog health.  The numbers of dogs within breeds has little to do with their vitality and much more to do with fashion and the whims of a handful of breeders.  No one would claim that a Pug is popular due to competitive natural gifts of robust health and fitness, rather they are much like the inbred wolves on Isle Royale, they are artificially supported by beneficial conditions, easy access to food, and a blunting of natural pressures against their survival; in the wolves’ case it’s the benefits of living on an island, in the case of the Pug it’s being coddled by owners and breeders willing to spend a pretty penny on their upkeep.

In open and competitive environments, population numbers can serve to estimate vitality when better data has not been taken, but it seems that on Isle Royale, the wolves have a sheltered niche as the apex predators with ample food supply.  The Moose can’t migrate away and there is little in the way of competition for the wolves.  In such conditions, even sickly inbred wolves can reach a carrying capacity at about the same numbers as more robust wolves can.  It’s possible that more wolves could have thrived had the main food source not plummeted, or it could be that even at the lowest levels the moose populations were not small enough to be a significant factor in the head count of the wolves.

Without a marked improvement in population size on the island and without documentation of the rates of disease and bone deformities improving with the influx of the immigrant wolf, the technical definition of a “genetic rescue” has not been met with the current state of knowledge about the Isle Royale wolves.

Claire, a disabled harlequin Great Dane on the first day of her new life after being rescued from a torture breeder where she suffered for 4 years. She is doing much better now.

Oh Claire,

I’m sorry for all the suffering you’ve experienced in your life.  It’s one thing for me to write about qualzucht, torture breeding, and to think and debate about it in the abstract.  But when theory hits reality, when real dogs with real names and real stories come forth, it’s nearly impossible to remain objective and dispassionate given the true suffering you qualzucht dogs go through.

You have been tortured.  You were genetically tortured by the breeder who created you when they deliberately bred two Harlequin Great Danes together, doubling up on at least two lethal genes (harlequin and merle) and possibly doubling down on other genes known to contribute to deafness and blindness.  It’s likely that more than half the puppies that shared your mother’s womb died before birth from these defects, but you survived.

You were further tortured by this breeder when they neglected to provide any real care for you, letting you suffer from a slew of debilitating conditions.  They allowed parasitic demodectic mange mites to run rampant and cover the majority of your body, they allowed parasitic yeast to invade most of your orifices, and they allowed parasitic worms to invade your heart.  Your hearing, vision, and immune system were trashed by your parasitic breeder leaving you defenseless against further exploitation.

You were tortured again when this breeder used you to produce Great Dane Puppies to sell in the open market and then dumped you in a shelter as soon as the puppies were sold.

Parasites are organisms that invade other life forms and leech sustenance at the expense of their host.  Mange, yeast, and heartworms are all lower life form parasites that prey on higher life forms.  The person who bred you is just a low life who has habitually tortured and exploited you and gave you nothing in return.

Man is the highest life form known to exist, yet the person who tortured you in so many ways is the lowest of the low.

I’m sorry Claire.  I’m sorry we did this to you by not speaking out stronger against horrible breeding practices and condemning parasite breeders.   I’m sorry that we rewarded your parasite breeder by buying your puppies.  I’m sorry that those breeders who consider themselves elite and the highest of the high do the exact same things that have caused you so much pain by breeding harlequin to harlequin, merle to merle.  They are parasites too.

But there are better people out there Claire and I’m glad you’ve finally found some at the Foster to Furever community, the people at Big Dog Rescue, and April Albin who has been fostering you during your recovery.  The last four years of abuse and starvation must have been hell, Claire, but you’ve been given a new life now.

So happy one month rebirthday, Claire, you’re already a new dog and on your way to a better life.

Claire, one month into her recovery. The mange can be treated but her compromised immune system is forever. Food can cure the starvation she suffered. But the deafness and blindness that were caused by her breeder are permanent and not treatable.

Cain said unto his brother, 'Come let us go out into the field' and when they were in the field , Cain set upon his brother Abel and killed him.

The Bible puts forth the killing of Abel by his brother Cain as the first murder in history, which foreshadows the eternal struggle between the farmer and the cowman.

Cain is the first man born of woman in the bible, and a second generation farmer–already lazy and living off the dole.  His goody-goody younger brother, Abel, was the first Sheeple in history and an insufferable brown noser.  Abel decided to one-up his older brother and gain favor with God by sacrificing one of his fat young lambs–mmmm baby lamb tastes good–making him the first enemy of PeTA.  Cain, not to be outdone had to make a sacrifice too, but since he was getting paid to not plant (as is the custom) he couldn’t really come up with a suitable sacrifice and he didn’t really want to burn those subsidy checks because all that fire would cause greenhouse gases to escape into the atmosphere.

Of course, God found favor in Abel’s sacrifice of a fat lamb and was unimpressed with Cain’s offering of a meager organic crop: eat that vegans, God eats baby lamb and thinks “organic” sucks.  Then God went all passive aggressive on Cain for not being more like his younger brother, so Cain kneed Able in the groin and beat his head in with the jawbone of that lamb Abel sacrificed: poetic justice for the evil baby lamb killer!  Thus, Cain is also the first eco-terrorist and murderer in history.

God, who totally fell for Abel’s ass kissing, was devastated and cursed Cain to never grow crops again (funny, we keep ‘cursing’ our farmers to do the same and they’re so pious they get all uppity when we even suggest that they can go back to doing their job and grow something) and like all trust fund kiddies, God suggested that Cain get a Europass and go backpacking in the East to find himselff.  But Cain wasn’t happy and told God that “they’ll totally hate me in the East” and God said “tell them you’re from Canada, they won’t know the difference.”

But since everyone likes the Canadians and since Cain was such a douchebag, God knew that the ruse wouldn’t cut it and so he placed upon Cain a mark such that anyone who saw it would know that they shouldn’t kick his ass. OR SO YOU WERE TOLD!

The truth is, if you look at the original Genesis passage, God didn’t place a mark upon Cain, he gave him a dog! At least according to one rabbinical scholar in the Midrash:

And the Lord set a sign for Cain.

R. Judah said: He caused the orb of the sun to shine on his account. Said R. Nehemiah to him: For that wretch He would cause the orb of the sun to shine! Rather, He caused leprosy to break out on him, as you read, And it shall come to pass, if they will not believe thee, neither hearken to the voice of the first sign, etc. (Ex. iv, 8). Rab said: He gave him a dog. Abba Jose said: He made a horn grow out of him. Rab said: He made him an example to murderers. R. Hanin said: He made him an example to penitents.

Rab’s interpretaion sounds sort of like Pit Bulls and Parolees, no?

See, taking that backpacking trip around the old country isn’t so entertaining when you’ve just killed the only other guy on the planet who isn’t your parents.  Without some young German chicks (since neither Germany nor chicks had been invented yet) staying in the same run down hostel to provide some companionship, Cain needed a good dog to keep him company in his forced solitude and to protect him.