Dog's Life: Lifestyle
Those little pests are nothing new!
Mummified dogs are not a new archaeological discovery, but finding bloodsucking parasites on them is. Over 400 dog mummies unearthed from the El Deir excavation site in Egypt have been found, and one young dog among them was infested with a number of parasites that have been preserved.
There were over 60 ticks found on this poor dog and there was one louse, too. The scientists who found this dog suspect that a tick-born disease that kills red blood cells was probably responsible for the death of this dog at such a young age. Besides the ticks and the louse, remains of two types of fly larvae were found on it, suggesting that the dog’s body had time to attract carrion flies prior to being mummified.
Mummifying animals was common in ancient Egypt. It was done to provide food and companionship for people in the afterlife and to make sacrifices to the gods, yet nobody is sure of the reasons for the dog mummies at El Deir. It is unclear if they had specific human guardians or how they died. Perhaps they were purposely bred to be sacrificed as cats commonly were, but we just don’t know.
Scientists involved with this excavating project are exploring questions about the source of the dogs. They are also hoping to find more parasites on the dog mummies in order to investigate the origin and spread of diseases and to deepen our understanding of the role of parasites in the history of the species.
Evidence that ancient dogs suffered from ticks, lice and other ectoparasites is prevalent in ancient writings such as those of Aristotle, Homer and Pliny the Elder, but this is the first archaeological evidence that corroborates those texts. It’s certainly no surprise that dogs living a couple of thousand years ago faced the danger and nuisance of ticks and lice. It would be astounding if it were a recent development in the lives of canids, but it’s still interesting to have such concrete evidence.
Culture: Science & History
Breeders, judges and historians talk about breed standards—why they work and when they don’t
In the world of mammals, the domestic dog— Canis lupus familiaris, a subspecies of the gray wolf— reigns as the most morphologically diverse. Consider, for example, the extremes represented by the 155- pound South Russian Ovcharka and the seven-pound Silky Terrier. This incredible variety can be attributed in part to the dog’s basic template, which can be customized by the manipulation of a very small number of genes.
For instance, 95 percent of all five canine fur textures and lengths (the Afghan Hound and Curly-Coated Retriever curiously excepted) is orchestrated by three genes. Further, just six or seven locations in the canine genome account for nearly 80 percent of dogs’ vast size and weight differences. (In humans, these genes number in the hundreds, if not thousands). A single mutation, shared by 14 diminutive breeds, determines that a dog will be small, and another is responsible for the long-bodied, short-legged nature of numerous dwarf breeds.
Clearly, the dog’s random morphology isn’t quite as arbitrary as we thought, and breeds aren’t quite as unique. Furthermore, canine traits come in packages. Flip a switch to make the legs more slender, and the skull will narrow as well. Turn down the volume on pigment and the chance of deafness increases.
Those who bred dogs had long known that traits were related, but there was little understanding of how those relationships worked; nor was there much concern. Dogs were bred for skills useful in a practical world. Once breed exhibition became a fashionable pastime and working dogs were awarded championships based strictly on appearance, however, all this changed.
In 1866, John Henry Walsh (writing under the pseudonym “Stonehenge”), editor of The Field, the most influential hunting and kennel journal in England, was the first to describe a breed’s physical characteristics with phrases that he believed were equivalent to its field ability. A bird dog judged perfect to a well-written breed standard would, by the logic of the day, perform perfectly in the field.
At the time, horsemen and sportsmen were the dog-fancy glitterati (women became active later), and many of the arcane descriptions in breed standards are borrowed from those arenas. For example, the Poodle’s “straight-forward springy trot” describes the dog’s ability to retrieve and carry a bird. The phrase “stand like a cleverly made hunter” references ideal anatomical construction and proportion in the German Shorthaired Pointer.
Today, breed standards serve three purposes: assessment in competition; delineation of unique qualities in different breeds, some very much alike; and maintenance of breed similarity throughout the world.
The question is, what happens to purebred dogs when language, intrinsically fluid and inexact, is used to suspend change in morphology and behavior? In the late 1990s, as a doctoral student in linguistics at Claremont Graduate University, I conducted a study to find out. Part of the research included interviews with experienced American Kennel Club (AKC) breeders, specialty judges and breed historians. What I heard from them provides some insight into specific ways that a standardized lexicon can influence change in pedigreed dogs far beyond what is intended.
Dogs in Translation For some breeds, international politics played a role. At the first Canine Congress in 1886, the Germans were opposed to the Swiss-type Saint Bernard, favoring the bulkier English type. Nothing was resolved until 1887, when the Swiss dog was finally approved as the international type. The United States club, with its strong ties to England, adopted the international standard in words, but in practice, bred to the English type.
During an interview, as three Saints gnawed on bones nearby, an experienced breeder and specialty judge offered his opinion: “This changed the morphology of the American Saint, most noticeably in the head. The Saint Bernard standard was translated, with some errors, from German to English in 1888. For instance, ‘when in action’ should have read ‘when excited or alert.’ The phrase, ‘the horizontal axis of the head’ should have read ‘the long axis of the head.’” More than a century later, the club had still not made corrections, perhaps because, as linguists argue, translation of a lexicon from one language to another can never be exact.
Translations are more like corrections or clarifications. When standards are clarified, they usually get longer and, consequently, more exclusive. A standard that calls for feet to be “round, compact, catlike, standing well upon the toe pads,” is more restrictive than one that says feet must be “close, round and firm.” Revisionists tread cautiously because an imprecisely rephrased standard can have an impact on a breed’s genetic diversity.
Amending a standard for any reason is controversial for those entrenched in a time-honored tradition devoted to blueblood history (albeit a fanciful history, since geneticists tell us that very few breeds are as old as they were once touted). As one Greyhound breeder observed, “The torch handed to us was the perfect coursing dog. Our standard is taken word for word from what Stonehenge wrote in the 1860s. If we added more words to make it more explicit, it may end up being a Greyhound different than the one each one of us has in our head.”
But progress necessitates change. A handful of words differentiate an apple, an orange or a pear, but 13 varieties of apples require a larger lexicon. In his 1576 treatise, Of Englishe Dogges: The Diversities, the Names, the Natures, and the Properties, cynologist John Caius described the generic land Spaniel in 58 words: “The most part of their skins are white and if they be marked with any spots, they are commonly red, and somewhat great therewithal, the hairs not growing in such thickness but that the mixture of them may easily be perceived. Other some of them be reddish and blackish, but of that sort there be but a few.” Today, Caius’s dog has morphed into 13 f lushing Spaniel breeds. The Field Spaniel standard uses 973 words, short in comparison to the English Springer Spaniel standard, a 2,040 word descriptor.
Occasionally, words are added to explain what something is not. As one breed standard committee member noted, “There were lots of questions from judges about the preferred shape of the eye opening. So we said it’s acceptable as long as it’s not this, that or the other thing.” And at a California dog show, pointing to Mastiffs benched only a few feet from his St. Bernards, an exhibitor told me, “Sometimes breed clubs have to lengthen standards to differentiate their breeds from others so similar that, if marked differently, could be shown as Saints.”
When Words Fail Dressed in formal attire appropriate for the straitlaced Madison Avenue cocktail party that precedes every Westminster dog show, AKC VIPs sipped their drinks and talked candidly about breed standards.
A Doberman breeder, specialty and all-breed judge opined, “I’m not sure that the standard hasn’t been what’s wrong with some of the breeds, in that by naming and describing the criteria with which the animal is to be judged, the words lack exactness. When people bred to the standard, the animal changed and became what the words described. Also, some groups wrote the standard and have not been able to breed to that ideal, so now, they change the nuance of the words to fit the breed ideal. Making the dogs fit the words, and not vice-versa, is wrong.”
For instance, a standard that establishes criteria to develop the best muzzle shouldn’t include terms that are subjective or indefinite. Or as one judge said, “If the standard calls for a short muzzle, judges select dogs based on the shortest muzzles in the ring. The breed’s muzzle gets shorter and shorter. So you have to ask, ‘Shorter than what?’”
Another said, “Our standard calls for the ear, when pulled forward, to reach the eye. You see so many dogs in the ring now with longer ears. We always say, well, which eye is it supposed to reach, and is it pulled under the muzzle or over it? I have never seen a Golden Retriever with ears too short.”
Some breeders get so fixated on one attribute of the traditional standard that they are willing to sacrifice something more important. “For example, they may create a broad head but are willing to accept shorter legs and a longer back in order to do so,” another judge observed.
Breed standards, like all nomenclature, are subject to the rules of language. Like the dog it describes, vocabulary is deceptively capricious and unexpectedly fluid. A good example is size. Big breeds are getting bigger. As I was told by a breed historian in reference to St. Bernards, “What was bred to be powerful and strong in 1900 would not be considered powerful and strong today. Like an automobile in 1915, it was powerful then but not compared to now.” If a standard describes a breed as strong and powerful, the ideal dog gets bigger.
Rather than hack away at standards, some breed clubs hold seminars for judges in which contemporary nuances of an indefinite vocabulary are refined. Others reluctantly reword phrases to accommodate inexperienced newcomers (currently, the average length of interest and activity in the dog fancy is five years or less). A Golden Retriever breeder told me, “People who wrote the original standard were horse people, and this is where the phrase, ‘deep through the heart’ came from. It had two meanings, deep through the chest and courageous. The original meaning and nuance of the old words is often lost on today’s breeders, or is interpreted to mean something else.” After years of debate, the phrase was reluctantly changed to “deep through the chest.” Some clubs don’t revise words, but instead, reinterpret their meanings. Take English Bulldogs, for example. As an owner of one of these stocky dogs remarked, “The interpretation of words has changed. The Victorian [Bull]dog was a transition dog, less bulky, less massive, taller, leaner, and is now thicker and more compact.” Another handler observed that “one of the issues in the standard is weight. It calls for 45 to 50 pounds. But it has no height restriction, so a higher-station dog might be thinner.” The 1910 dog was a much taller and leaner dog compared to today’s stout fireplug variety, but both are considered to have been bred correctly to the standard.
A German Shepherd breeder and specialty judge who chairs the club’s standard committee told me that “you can have the same words in several standards, but they don’t mean the same thing. We use the word ‘almondshaped’ in our standard. But if you look at other breed standards, both the Collie and American Cocker call for almond-shaped eyes. The Collie has a small triangular eye and the Cocker’s is a goggle-eye [the eye protrudes from the skull].”
Judges Play a Role A specialty judge is an experienced breeder and expert on a particular breed. An all-breed judge is a generalist qualified to judge several breeds. The specialty judge brings meaning to the words in the standard, and the allbreed judge makes sure the words mean what they say.
A specialty judge who also works as an all-breed judge explained that the interpretation of complex descriptions, such as the angle of the hock, is more difficult for a generalist all-breed judge to measure. “Because it’s easier to see a proper bite than a proper angulation, the bite may be given more significance than something more important, such as angulation of the hindquarters.” On the other hand, the all-breed judge tests the words. “If the breed club thinks the all-breed judge is misinterpreting the standard, then they need to rewrite it. The judge shouldn’t choose the dog that he thinks they mean.”
War of Words The AKC considers itself a club of clubs. Owners intent on breed registration must first demonstrate that a majority of breeders are interested in establishing a national breed club. Who gets to be in that club is at the heart of a mounting number of controversies.
In 1994, the AKC Labrador Retriever standard was revised to exclude dogs less than 22 inches at the withers (or 21 inches for bitches). Some breeders whose dogs no longer met the standard were part of an $11 million class-action suit against the AKC Labrador Retriever Parent Club (the national organization designated by AKC to represent the breed), claiming that height restrictions excluding shorter dogs no longer described the Labrador Retriever: if you make a bigger dog, you make a different dog. A litigant told me, “It’s perfectly reasonable to change a breed, but the dog should have a different name.” They tried and failed to trademark the name Labrador Retriever; the judge sided with the AKC parent club. The Border Collie war began in 1988, when the American Border Collie Association and others heard rumblings that some wanted to register the breed for conformation showing, which requires a breed standard. This idea didn’t go over well with herding trial enthusiasts; a Border Collie is what it does, not what it looks like. Any dog can enter an open sheepdog trial. There are no age, size, color, shape or breed restrictions, and registration is not required. Unlike registered purebreds, whose lineage must be proven in ancient studbooks, many Border Collie champions are registered on merit (ROM). In theory, a Pomeranian who could prove its worth at a sheepdog trial could, by performance, be called a Border Collie.
In the minds of many, AKC conformation specifications threatened 200 years of breeding for performance, not looks. Led by Donald McCaig, who retold the tale in his book The Dog Wars (2007, Outrun Press), the group prepared for battle: “Hands off the Border Collie! We own Border Collies. Our dogs are companion dogs, obedience dogs and livestock-herding dogs. For hundreds of years, Border Collies have been bred to strict performance standards and today they’re the soundest, most trainable dogs in the world. The AKC wants to push them out of the Miscellaneous Class and into the show ring. They seek a conformation standard [appearance standard] for the breed. We, the officers of every single legitimate national, regional and state Border Collie association, reject conformation breeding. Too often, the show ring fattens the puppy mills and creates unsound dogs. We will not permit the AKC to ruin our dogs.”
They filed to legally trademark the name but, like the Lab litigants, lost in court. In 1997, the first Border Collie was shown in conformation at Westminster. I was there that year and interviewed a handler/owner who had been instrumental in getting the breed registered and in writing the standard. I asked her how she did her research. “This dog is shown in Australia, the British Isles and New Zealand. So I read their standards and asked them what they would do differently if they could. I tried to emphasize movement and gait. The standard shouldn’t describe a still dog. The Border Collie is almost a vision of movement even when it is standing still … always poised on the brink of action. The head drops for a reason. It is common knowledge among Border Collie people. That’s why I didn’t include a description of the head in the standard. I thought everybody would know that.”
In the benching area, surrounded by panting dogs crowded into crates and standing patiently while being primped on grooming tables, the woman sat in a folding chair, visibly distraught. Tearyeyed, she continued: “Right before you got here, a prominent breeder came by and said he will breed a dog with the head held higher. The head has to do with movement in the field, in making eye contact with the sheep, the pattern of behavior that has evolved from two centuries of work. Now I regret having fought so hard for this. The standard should not threaten the dog as a working animal, but I believe it now will.” About the same time the Border Collie war was raging, AKC enthusiasts saw an opportunity to register the Jack Russell Terrier (JRT), an irascible, independent dog with an intense work ethic, extremely diverse genome and phenotype as dissimilar as that of the Border Collie. Many Jack Russell Terrier breeders vehemently opposed the action, claiming that the breed’s physical and working characteristics would be jeopardized by this move. Nevertheless, the splinter group formed the requisite national breed club, named itself the Jack Russell Terrier Breeders Association (JRTBA) and gained AKC recognition in 2001.
A lawsuit ensued. After an expensive court battle, the name Jack Russell Terrier was awarded to the working phenotype and the AKC changed the conformation dog’s name to Parson Russell Terrier. The AKC parent club is now the Parson Russell Terrier Association of America.
Writer Alston Chase, who includes the story of the breakup of the breed in his book We Give Our Hearts to Dogs to Tear (2008, Transaction Publishers), told me, “The Jack Russell Terrier is a feisty, very aggressive, very tough dog. But middle-class urbanites don’t really want that kind of dog. They want a dog that will be a good pet.” Chase, who lives in rural Montana, has bred the working terrier since the 1970s.
The public took notice of the rather obscure breed when it became a media darling in the 1980s. Chase said, “The overbreeding followed the popularity driven by the media, not by the dog itself.” Aggressive and difficult in a pet environment, the breed was misrepresented as a mischievous lap dog on shows like the NBC sitcom Frasier. Surprised and disappointed by their dogs’ ornery personalities and exercise requirements, urban pet owners abandoned JRTs at shelters in record numbers.
According to Chase, one of only a handful of people in the U.S. continuing to breed the old-fashioned dog, “We’re doing what we can to prevent extinction of the original breed, but people aren’t in love with the value of diversity in the dog. They want dogs that look alike.”
Geneticist Jasper Rine, in a letter to the AKC supporting the Border Collie anti-conformation campaign (included in the appendix of McCaig’s 2007 book), predicted what was to come. “It may be nearly impossible to breed for a particular behavior based on heterozygous advantage and still achieve a homogenous conformation.” Breeding dogs for fixed conformation means breeding for homozygosis (the formation of genetically identical gametes) of the genes that contribute to appearance. In doing so, genetic linkage (the tendency of genes located in proximity to each other on a chromosome to be inherited together during meiosis, or cell division) may result in genes near those controlling conformation becoming homozygous as well. Unfortunately, chance determines which genes are swept up. By breeding for conformation, breeders may be breeding away from desirable behavior, even putting alleles (forms of a gene) at risk for extinction.
So, what has happened to the Labrador Retriever, Border Collie and Jack Russell Terrier over the last 15 years?
The shorter-legged, more compact field-bred Lab continues to be shown in Canada, the UK and other countries that don’t disqualify individuals based on size. In the U.S., conformation and companion Labs are getting increasingly larger. Will diseases linked to large size compromise the American line? Time will tell.
As a consequence of the acrimonious Border Collie war, few working dog breeders had a desire to become specialty judges, so the fate of the conformation dog was left in the hands of generalist judges who lacked sheepdogtrial experience. As predicted, the standard created a split type: working dogs continue to be a rag-tag group, dissimilar in shape, size and color, but the same in their relentless determination to move sheep from one place to another. In contrast, AKC dogs look very similar, but their ability to herd sheep is open to question. Are both types called Border Collies? Formally, yes, but the AKC dog is widely, popularly and even affectionately known as the “Barbie Collie” by some: pretty as a picture, but, according to the member-owned American Border Collie Association and others, as blandly attractive and vacuous as the doll from which the name derives.
The working Border Collie is safe for now, but the old-fashioned Jack Russell Terrier can hardly be found. Like the Old English Bulldog who faded away with the passage of the 1835 Cruelty to Animals Act and the Wolfhound who died out with the demise of the wolf in the British Isles, the pre-AKC JRT will likely disappear as well. Eventually, sentimental breeders may attempt to recreate the breed when they realize what they’ve lost, and they may perhaps have some success in replicating the way the dog appeared. But the breed’s signature obstreperous temperament is something people will only read about in books.
* * * * *
It would seem, then, that words are indeed powerful. To say that the lexicon used to describe a purebred dog, or even name one, will not affect the way we engineer the animal contradicts the language-relativity hypothesis, which holds that the vernacular we use to frame our perceptions influences the way we regard, understand, interpret and reinvent them. As a tool, language plays an important role by which innovation—in this case, of a sentient human-made domestic animal—is further developed. Or, as the AKC says, refined.
To paraphrase Shakespeare, would a rose by any other name smell as sweet? If you’re talking about dog breeds, that’s not a rhetorical question. The answer is no.
Dog's Life: Lifestyle
Conflict among scientists who study it
Research about canine genetics and the domestication of dogs is an exciting area of study with many players, so it should surprise nobody that there is disagreement within the field. Multiple groups of researchers from around the world have compared the genomes of dogs and wolves. While they generally agree about the genetic changes that have produced differences between dogs and wolves, their conclusions about the domestication of dogs vary wildly.
The disagreement concerns fundamental aspects of the evolution of dogs such as where, when and why dogs evolved from wolves. So, the location, the timing, and the reason for domestication that various groups propose are not even close.
One group suggests domestication occurred around 10,000 years ago in the Middle East and that it was the development of agriculture around that time that was the catalyst for domestication. Another group claims that it happened around 32,000 years ago in the south of China and related to scavenging alongside the people living there. A third group narrows the time frame for domestication to between 16,000 and 11,000 years ago, and believes that the wolf population from which dogs arose is extinct, making it hard to determine the location of domestication. This third group believes that dogs became domesticated near hunter-gatherers rather than in the presence of an agrarian society.
Much has been made about the discord among scientists studying the domestication of dogs, but it’s hardly surprising. The cutting edge of science is always marked by strongly held opposing views. In the best situations, the intense disagreement among people working in the same field is a crucial part of making progress. Competing hypotheses are critical for the advancement of science. As people challenge each other’s views, all are spurred to study the subject more deeply and design experiments to investigate that which has been called into question. From the ongoing work, the conflicts are eventually resolved as some ideas fall by the wayside and others gain increased support from new data and discoveries.
Sometimes the conflict is cordial and in other cases, it can be very bitter. At this point, the scientists studying dog domestication say that though there is a certain amount of rivalry, they get along and enjoy talking with each other. That may be harder to maintain as people move to the next phase of research into dog domestication and seek to sequence DNA samples from ancient dogs and wolves. The availability of archaeological bone samples is extremely limited so there will be a lot of competition among scientists for both funding to conduct the research and access to the material necessary to do so.
In other words, we can expect a lot of fights over bones in the near future.
Celebrating the elegant life … with dogs.
Like many 19th-century painters of modern life, French artist James Tissot (1836–1902) frequently depicted the new, more intimate relationships between dogs and their owners. During this period, people increasingly believed that animals and humans had similar emotional and intellectual responses, and the bond between pet-keepers and pets was foregrounded in new ways. Tissot, an avant-garde painter associated with Edgar Degas, portrayed a wide range of dog types with great charm, affectionate understanding and skill throughout his career. Indeed, he must have owned a number of dogs, as particular animals appear repeatedly in his pictures. A painting from Tissot’s French period, Young Lady in a Boat (1870), represents a woman in the fashionable costume of the early 19th century, accompanied by a Pug. This toy breed was a sign of wealth and status. Here, it has the slightly longer legs and snout that were typical until later in the 19th century, when the dogs began being bred for the pushed-in faces popular today. The woman’s gaze suggests that she is on a romantic rendezvous, and indeed, an alternative title for the picture—Adrift—implies that she could potentially lose her moral compass as well. Clearly, her Pug is keeping a watchful eye on her!
Culture: Science & History
Village dogs’ genetic code may hold clues to canine evolution and health
Like classic twin studies that investigate the interplay of nature and nurture, comparing the genome of village dogs to modern dogs may help disentangle the long-term evolutionary effects of genetic and environmental influences.
Mastiff to Min-Pin, Corgi to street cur: all dogs share the same set of roughly 20,000 genes. What makes one dog different from another—or, in the case of purebreds, almost the same— is how the genes are expressed and restricted from being expressed, and how they communicate with one another. Therefore, it may be safe to say that each of the world’s 800 to 900 million dogs is a distinct combination of different versions of the same genes. Or maybe not. At least, that’s what some scientists suspect, and they think they’ll find answers in the DNA of the ubiquitous, free-ranging canine outcasts that populate developing countries throughout the world.
While village dogs were being socially shunned, modern dogs—a subpopulation that likely split off from village dogs thousands of years ago—were serving society. So tightly woven into the fabric of our lives that we rarely think of them as human-engineered, dogs have been refined for increasingly specialized tasks such as hunting, transportation, protection, warfare, ornament and companionship. As a result of rigorous artificial selection over a long period of time, many of their ancestral gene variants are suppressed. Some have disappeared altogether, creating a fragile homozygous genome that has little diversity.
In contrast, village dogs are barely tolerated by society. Although considered a domestic species, they are the products of thousands of years of natural selection. Consequently, their heterozygous genomes are robust and extremely diverse. In addition, it’s possible that long after modern dogs branched off from the family tree, some village dog populations may have developed new gene variants that protect their immune systems.
Evolutionary biologist Adam Boyko, assistant professor in the Department of Biomedical Sciences at the Cornell University College of Veterinary Medicine, is confident that comparing and contrasting the two branches of the domestic canine family tree will provide answers to some of the mysteries that continue to surround the evolution of the domestic dog: When and where were dogs domesticated? What were the global migration paths of humans and dogs? What genetic changes occurred when wolves became dogs? Which genes are responsible for extreme size, shape and behavior differences? What are the underlying causes of genetic diseases? And how do parasites have an impact on canine well-being?
As a postdoctoral student at Cornell, Boyko worked under the tutelage of Carlos Bustamante, now professor of genetics at the Stanford School of Medicine. Curious about how the underappreciated and even less-studied village dog genome might reframe our current understanding of canine evolution and domestication, Boyko and Bustamante persuaded Ryan and Cori Boyko (Boyko’s brother and sister-in-law, who were then both graduate students in anthropology at the University of California, Davis) to add a few side trips to their otherwise romantic African honeymoon. Their instructions were to catch semi-feral, uncooperative village dogs and draw blood samples, then ship the samples back to the lab for analysis. Information from the preliminary DNA samples indicate that the researchers are on the right track. I asked Dr. Boyko about his research, and if it has future application to invigorating the health of our companion dogs.
Jane Brackman: How will mapping the genome of the village dog help us understand the mechanisms of traits in modern dog breeds?
Adam Boyko: Geneticists have spent a lot of time looking at purebred dogs. When something is selected for, either by natural or artificial selection in a population, geneticists can tell because of the patterns that are left in the genomes of individuals in those populations. In humans, for example, we can clearly see that lactase persistence, the ability to digest milk into adulthood, was selected for in some populations.
When we look for these patterns in purebred dogs, we find that things like ear f loppiness and tail curliness are driving these patterns, or short legs or small/big size. Basically, we find the effects of artificial selection by humans for breed standards. If we did a similar scan for selection in village dogs, perhaps some of those same genes would show patterns of selection, but I think we’d also see a new class of genes showing patterns due to natural selection.
For example, maybe there was a lot of selection in early dogs for genes in certain metabolic pathways because there was such an extreme dietary shift from wolves to dogs. Or maybe new parasites and pathogens caused selection at genes influencing the immune system. Or maybe we’ll see selection around genes that influence behavior and temperament.
Basically, there are all sorts of theories about how dogs became domesticated and what makes a dog a dog. When we look at purebred dogs, the main thing we are able to see is what makes certain dog breeds look and behave one way versus another. Maybe by looking at village dogs, which are much less influenced by the strong and recent artificial selection taking place in breed dogs, we’ll be able to see patterns of selection that occurred earlier in dog history.
JB: Might your findings have application for the future? For example, if you were to come across genes influencing the immune system, would breeders be able to use that information to revitalize the pedigreed-dog immune system?
AB: It is certainly true that my research may find a new MHC-type immunity gene [the major histocompatibility complex mediates the immune system’s white blood cells] that has been lost in many purebred dogs and which could reinvigorate their immune diversity. Or perhaps it will find variants associated with diet, and make us start considering a dog’s genetic makeup when making dietary recommendations. But I’m really not comfortable speculating, since I’m likely to be quite wrong in these predictions.
For example, I would have never guessed that deliberately infecting patients with intestinal parasites [Helminthic therapy] would cure ulcerative colitis, but that seems to be the case, and signatures of selection in the human genome help explain why.
But having said that, I think looking at the genomes of village dogs will be extremely useful. For example, we could get a better picture of the kinds of traits that were selected for in natural dog populations, including disease resistance, which might give us useful insights into diseases we diagnose in our pet dogs.
Conversely, as veterinarians and geneticists find more mutations that cause disease or unique traits in dogs, we can look at the genomes of diverse village dogs to see when and where these mutations arose, and whether they are also found in any other village or purebred dog populations.
It’s a really exciting time to be a canine geneticist, as we have all these new genetic tools at our disposal and many, many purebred and free-ranging populations that have yet to be characterized genetically.
JB: Some populations of village dogs, such as those you’re studying, have been isolated for many thousands of years, evolving under pressures that the stem parents of modern breeds were never exposed to. Is it possible that these dogs have “new” gene variants that don’t exist in the genome of modern breeds?
AB: It’s certainly possible, and it’s something I’m very interested in. For example, my lab is looking at free-ranging dogs in the highlands of Peru to see if they have any genetic adaptations for high altitude. Perhaps more importantly, some village-dog populations might harbor disease-resistant variants for parasites or pathogens that are prevalent in their area, but these variants might not have made it into modern purebred dogs, since those breeds were mostly founded elsewhere.
JB: How urgent is it that we learn more about these ancient dog genomes?
AB: We know how quickly pre-Columbian Native American breeds were lost when Europeans brought dogs with them to the New World, and we see that it will happen like that very soon in other remote parts of the world. So we’re working as fast as we can to get the data before the dogs are gone.
JB: What’s going on in your lab now?
AB: We’re collecting DNA samples and the genetic information we need so we can start piecing together what’s going on in these interesting but largely neglected free-ranging dog populations. We are seeking insights into dog population history to discover patterns of selection around certain genes that can then become the basis of further study. Our work is very hypothesis-driven. We have certain hypotheses about how dogs evolved, and we try to collect the right samples to test these hypotheses.
As geneticists learn more about how genetic variation controls complex traits in purebred dogs, we find it’s quite different than what we see in humans. Why? There are at least two competing hypotheses, and by gathering data from free-ranging dogs, we can start testing them to figure it out. Some of this gets into technical discussion about genetic architecture, recombination, epistasis and pleiotropy and such, so I’ve avoided getting too academic. But I also don’t want to be dismissive of it since those technical, hypothesis-driven aspects of the project are the bread and butter of my lab in terms of student training.
JB: In longitudinal studies such as the Morris Animal Foundation’s Golden Retriever Lifetime Study, researchers gather information from participants’ DNA and then match what they find to traits the test dogs may display over a lifetime. Will you have an opportunity to see how, for example, an immunesystem mutation affects a village dog’s health as it ages?
AB: Our project is a huge undertaking, and there’s a ton of data we’d love to gather on each dog but just simply aren’t able to since, at this point, we’re focused on sampling as many dogs from as many populations as possible to maximize the amount of diversity we can analyze. I really don’t want to overstate what we’re able to do in one visit to check out a dog and draw blood, which is limited to looking for genetic signatures in the genome of these dogs showing signs of selection and/or local adaptation.
But, since we have a fairly good idea of what genes do in modern dogs, at least in a rough sense, if we see a genetic signature in village dogs for positive selection around a gene we know is involved in immune function (for example), that’s a big discovery.
JB: At the risk of oversimplifying, say you’re looking at a region that you know to be linked to a negative trait and you see that the switch is turned off in the village dog DNA and turned on in modern dog DNA—would you feel that you’d found a “smoking gun”?
AB: It’s possible. Then, of course, as you allude, we would want to go back, look at dogs carrying that mutation versus other dogs, and see if there are different health outcomes. Perhaps [dogs with the mutation] are more resistant to intestinal parasites or perhaps they are more prone to autoimmune disease or something. Until we find the mutations, it’s a bit speculative to make predications about what exactly the findings will mean to owners. This is certainly “basic research” in the purest sense.
JB: In people, size is determined by hundreds of genes, each with a small effect. In purebred dogs, body size can be regulated by a single gene. Is this unique to dogs?
AB: It depends. There are other traits in other species controlled by a couple of loci [location of a gene on a chromosome]. I would argue that yes, it’s pretty unique. Whether or not dogs are special in that there is something about their genome that predisposes them to this type of diversity, or perhaps because humans worked so hard at creating them, we don’t know. This debate is still raging in the literature. It is definitely the case that genes have many, many effects. Rather than being a blueprint in which each gene is responsible for just one part of building the whole organism, the genome is more complicated, with each gene taking on different roles at different times or in different tissues.
JB: Do multiple-trait relationships also show up in village dogs?
AB: I think this would also occur in village dogs if the mutations were in those populations. The difference is that selective breeding has actively promoted these large-effect, diversifying mutations in dog breeds, making them relatively more common. Natural selection usually selects against such large-effect mutations in natural populations. You won’t see a short-legged wolf because it couldn’t hunt.
In fact, most of these large-effect mutations probably first arose in village dogs. The difference is that these mutations aren’t usually beneficial to village dogs, but the ones that aren’t too detrimental might persist at low frequency long enough for humans to start trying to promote breeding of that trait. Take achondrodysplasia [a type of dwarfism]. It almost certainly arose in village dogs, but to a free-ranging dog, super-short legs and all that comes with them probably aren’t much of a selective advantage. But once folks started looking for dogs to turn their spits, they found these super-short dogs to be useful, and eventually that genetic variant made its way into a whole host of modern breeds.
For the specific achondrodysplasia mutation, I don’t know if that is the exact story, but I do think this is likely to be the case for many large-effect mutations. Depending on how early in dog evolutionary history the mutation arose, it could be found in most regional village dog populations, or it could be restricted to certain populations that are close to where it first arose. Lots of research still left to be done!
JB: As a lifelong dog lover, you must find it difficult to see the deplorable conditions in which some of these dogs live.
AB: There’s so much disease in these high-density populations. As these communities become more urbanized, dogs are living like rats and pigeons. Getting DNA on these populations is not enough of a reason to allow the animals to exist like this. Life on an urban street is rough existence.
JB: If you adopt a village-dog puppy and raise it in a typical Western environment, what kind of dog will you have?
AB: Adopting the dogs is not part of our project, but we know people who have done this. They can be great dogs. They don’t have some of the aggression issues you might see in some of our dogs, because they are culled for aggression, or for eating a chicken. There are some things that aren’t tolerated. So you might say that people in the villages impose a form of selection. The dogs are smart and resourceful. They seem to adapt.
Culture: Science & History
"Though in their deep heart’s core, there is a commonality of origin, spirit, emotional intelligence and empathetic sensibility, the wild wolf looks through us, while the dog looks to us. "
Of all the myriad members of the animal kingdom, the domesticated dog (Canis lupus familiaris) is closest to us. With individual exceptions in other species, this canine species is the most understanding, if not also the most observant, of human behavior—of our actions and intentions. This is why dogs are so responsive to us, even mirroring or mimicking our behavior. And it is why dogs are so trainable.
Fear in unsocialized and abused dogs interferes with their attentiveness to and interpretation of human behavior and intentions. This is one reason wild species like the coyote and wolf, even when born and raised in captivity, are difficult to train. The wolf “Tiny,” whom I bottle-raised and intensely socialized during her formative early days, never really lost her fear and distrust of strangers.
Tiny did not start mirroring human behavior until she was close to nine years old. At this point, she began to mimic the human-to-human greeting grin, revealing her front teeth as she curled her lips into a snarly smile. In my experience, dogs who can do this do so at a much earlier age, even as early as four to six months.
In comparing socialized (human-bonded) wolves and dogs in terms of how they have related to me as well as to my family members, friends and strangers, I would say that the main difference between the two species is the fear factor. Differences in trainability hinge on this; as I theorize in my new book (Dog Body, Dog Mind), domestication has altered the tuning of the dog’s adrenal and autonomic nervous systems. This tuning (which dampens adrenal fright, flight and fight reactions and possibly alters brain serotonin levels), is accomplished through selective breeding for docility, and by gentle handling during the critical period for socialization.
According to the earlier research of my mentors—Drs. John Paul Scott and John L. Fuller of the Jackson Laboratory in Bar Harbor, Maine—pups with no human contact during this critical socialization period (which ends around 12 to 16 weeks of age) are wild and unapproachable.
When we peel off the wolf’s innate fearfulness and put it on the dog, we turn the dog into a feral facsimile of a wolf. But a human-socialized wolf without fear could be an extremely dangerous animal, even attacking a human perceived as a pack rival. This happened to me in the 1970s during the filming of the NBC documentary, “The Wolf Men.” An alpha male wolf, along with his female cagemate who was in heat, had been released into a large wooded compound belonging to my friend, the late wildlife illustrator and conservationist Dick Grossenheider. Earlier that memorable morning, the she-wolf had greeted and solicited me, a total stranger, when I had visited the two wolves in their enclosure. (I am not saying that a male dog would never react to me in a similar way under comparable circumstances; I was once urinated on by the alpha male lead sled dog of a well-known racing pack in a similar situation.)
Canine Evolution and Human Needs
We see many things in our dogs’ eyes, the windows of their souls. They are also mirrors of the human soul, since every pair of dog’s eyes reflects—for better and for worse—how well that dog has been treated by our own kind. Dogs read our eyes and are attentive ethologists of human behavior, action, emotion and intuition. A change in tone of voice can make a dog tremble in fear or dance and yap for joy. Such ability to read human behavior, intentions and emotions was naturally selected for as dogs domesticated themselves and adapted to life with Homo sapiens, the “killer ape.”
Civilizing the Killer Ape
These half-human killer apes were never to be fully trusted, however, since when they were very hungry, or warring between themselves, they would often engage in cannibalism, as today, people still kill and eat dogs for sustenance and “good medicine.” (See James George Frazer’s The Golden Bough.).
Just as not every dog is fully domesticated, in that some can still turn feral, and others turn on their masters, so the wolf has failed to fully domesticate and civilize the killer ape. We still wage war with our own kind, and ignore the biological wisdom and prescience of such injunctions as do no harm, resist evil and treat others as we would have them treat us. All good dogs know this, and show it in their eyes and behavior. (But they have not forgotten how and where to bite!)
The secure, well-loved and understood dog is more often an extrovert than an ambivert wary of strangers. Secure, well-loved and understood wolves, in contrast (with few exceptions, some of whom become more easy-going around new people and in new places as they grow older, as did Tiny in her early teens), are more often ambiverts or introverts. They are fearful when meeting unfamiliar people. Differences in individual, breed and species autonomic tuning (as mentioned earlier) also account for differences in disease resistance, temperament and learning ability.
The Canid Conscience
Good dogs can see and respond to our own deep heart’s core of love and devotion because it is from this center of our own being that we embrace and celebrate theirs. That is what Franz Kafka in his essay, Investigation of a Dog, meant, I believe, when he wrote: “All knowledge, the totality of all questions and all awareness, is contained in the dog.” And this is why the ancient Egyptians believed that dogs were our guide in the afterlife—they were such good guides and loyal companions in real life. Embodying finer qualities of feeling and sensibility than the relatively irresponsible and emotionally challenged average human, dogs are worthy of being looked up to with awe and gratitude. We should help others of our own kind feel and know that in the deep heart’s core of all good dogs and wild wolves lies the source of an abiding affection that we, in moments of grace and communion, may share.
Since this core is as evident in a wolf as it is in a Toy Poodle, it is clear that neither domestication nor wildness has altered their true natures. In the heart of every dog is the spirit of the wolf that embodies the finer qualities of human nature that we call love and devotion.
There is a really interesting article posted on The Atlantic site today about the popularity of large dogs in China. As the author Damien Ma notes, “Most Americans will likely have a preconceived notion of the Chinese relationship with dogs. When a developing country can barely take care of all its own people, animal rights tend to sit very low on the totem pole. But the reality is much more complicated, especially with a burgeoning dog culture associated with the rise of young urban elites with disposable income. “
Ma then interviews an American filmmaker who is making a film Oversized Dogs: Chinese Dog Laws and the People Who Break Them. The director, who remains nameless for now, has been interviewing Chinese dog lovers who, similar to many dog lovers in other countries, find laws pertaining their pets onerous at worst, and turning many into scofflaws . But this isn’t a simple examination of a rising middle class pleasure in having dogs and their attachment to pets, it really does say more about how societal attitudes in China are evolving. As the director remarks: “From this, I realized that Chinese individuals casually break laws everyday, and this constitutes a very subtle and interesting side of dissent.” Read the whole interview and find out more about this Chinese “secret dog society” and what it might portend for the future of dogs in the world’s most populous country.
News: Guest Posts
I had watched the dog origin wars as a chronicler of the dog-human relationship for several decades when in 2009 I was approached a young editor The Overlook Press about writing a book on the origins of the dog. I readily agreed, and the result was How the Dog Became the Dog.
Pondering the conflicting dates, places, and theories associated with the emergence of the dog, I concluded that as soon as our forebears met wolves on the trail they formed an alliance of kindred spirits, and the process began. Their basic social unit was a family with ma and pa at the head and young ones of varying competency. They worked and hunted cooperatively. They were consummately social but capable of prolonged solo journeys.
It made sense that the Middle East, if not North Africa, was where this all started because that would have been the region of first contact. But because of their natural affinity, wolves and humans got together wherever they met. Some of the resultant “dogwolves”—my phrase for doglike wolves or wolves that act like dogs—created lineages that survived a while then fizzled out; others endured.
I identified several hotspots for early dogs across Eurasia and a group of humans that at least according to genetic evidence might have made its way through the cold of the last Ice Age from the Persian Gulf oasis, then a fertile land, to the Altai Mountains of Central Asia, a region that also hosts the headwaters of the Amur River, still famous for its wildlife. This group’s dogwolves mixed and matched with others along the way, especially the big mountain dogs of the Caucasus. This group of hunters and foragers gathered in the Altai around 40,000 years ago and from there ultimately took the New World.* They also went with their dogs, I calculated, south and east into China, Korea, and Japan and west again with their giant dogs, now mastiffs.
I based that conclusion in part on the types of dogs found in the New World. It made more sense that the possibility for the phenotype was present even if the phenotype itself was not manifest than that it was introduced later.
It was with some interest, then, that I read in PLoS One for July 28, 2011, about a 33,000 year old ‘incipient” dog from the Altai Mountains—that is, an early attempt at a dog that went nowhere. The finding was immediately challenged, and the fossil dismissed as a wolf, even if a strange one. So a new team of researchers redid the work in Robert K. Wayne’s evolutionary biology lab at UCLA and on March 7, 2013, published an article in PLoS One confirming that the 33,000 year-old-fossil is that of a primitive dog.
Writing for their colleagues from Russia, Spain, and the U.S., Anna S. Druzhkova of the Siberian Branch of the Russian Academy of Sciences and Olaf Thalmann of Turku University, Finland, state that when compared with other canids, the Altai dog, as it is known, shows closest affinities with New World dogs and modern dog breeds, ranging from Newfoundlands to Chinese Cresteds and including cocker spaniels, Tibetan mastiffs, and Siberian huskies.
Equally interesting from my perspective, the Altai dog does not appear to have been related closely to wolves in its immediate vicinity or to modern wolves. It came to the Altai from elsewhere, probably with people.
The researchers emphasize that there is uncertainty in their findings because they are based on a single region of mitochondrial DNA. But from my standpoint, the work provides one bit of evidence that’s I’ve not been barking up the wrong tree—and that seems worth noting.
*Ted Goebel et al., “The Late Pleistocene Dispersal of Modern Humans to the Americas,” Science, March 14, 2008. Connie J. Kolman et al., “Mitochondrial DNA Analysis of Mongolian Populations and Implications for the Origin of New World Founders,” Genetics, April 1996.
Culture: Science & History
Which DNA test should you choose to settle the “what’s in the mix” question?
For years, you’ve argued with your spouse that the 60-pound, black-and-tan tennis ball–chaser who takes you for walks and sweeps the coffee table clean with his tail is a German Shepherd mix, and that there is absolutely no Doberman Pinscher in there. Finally, in order to end the breed debate once and for all and restore peace to your household, you’ve decided to settle the question with a mixed-breed analysis test.
You’ve heard of the “swab test” and the “blood test” and know that both claim to unravel breed ancestry. With a little more research, you discover that the world of canine heritage tests has expanded since the first tests became available in 2007. Having a choice is great, but how do you go about comparing them and choosing the one that’s right for you?
There are several factors to consider, including the type of sample required, the number of breeds that can be identified, costs, turnaround times and the way the results are reported. Before you commit to a test that will decide the outcome of the German Shepherd/Doberman Pinscher battle, be sure you understand what you’ll be getting.
MetaMorphix Inc. (MMI) Genomics administers the Canine HeritageTM Breed Test, commonly referred to as “the swab test,” and Mars Veterinary provides “the blood test,” the Wisdom PanelTM MX Mixed Breed Analysis. These two companies have been considered the main players in this market, but new contenders are flocking to the scene. The most recent challengers are DNA Print Genomics, which offers the Doggie DNA Print, and BioPet Vet Lab, which recently unveiled the Dog DNA Breed Identification Test. Both use cheek swab samples.
The swab sample has the advantage of a collection procedure that is simple enough to be done by the owner at home. It does have some drawbacks, however, including a risk of contamination and too few cells being obtained for successful testing. To avoid the latter, BioPet Vet Lab includes a card that changes color to indicate that a sufficient sample is present. Blood samples are collected by a veterinarian and the chances of contamination and inadequate sample size are greatly reduced.
Tests also differ in the number of breeds available for comparison. Mars Veterinary interrogates the genetic signatures of more than 130 of the 159 breeds recognized by the American Kennel Club (AKC).* MMI Genomics recently announced the XL version of their test, which has a database of more than 100 breeds. The breed list available for BioPet Vet Lab contains 58 breeds. DNA Print Genomics does not report specific breed matches; rather, “15 elements of dog ancestry” are revealed, and the customer performs a search of the company’s online database to identify matches to particular breeds. Since each test interrogates a different set of breeds, sending your dog’s sample to more than one company may not return identical results.
Cost and turnaround times vary from test to test. In the past, the cost of the Wisdom Panel MX Mixed Breed Analysis test was determined solely by individual veterinary clinics, but the test can now be purchased online for $124.99, and test results are available within three to four weeks. (Your veterinarian will still have to draw the blood sample.) The swab tests are sent directly to the owners for sample collection, and prices and turnaround times vary: Doggie DNA Print, $199, six to nine weeks; Canine Heritage XL Breed Test, $119.95, four weeks (if you submitted a sample for the original version of the Canine Heritage Breed Test, you can purchase an upgrade to the XL version for $55 online); and the Dog DNA Breed Identification Test, $57.95 to $59.99, two weeks. (Prices current at press time.)
Results are presented as a certificate or report, depending on the company. MMI Genomics provides owners with a Certificate of DNA Breed Analysis. Three breed categories are included in the results: Primary, Secondary and In the Mix. BioPet Vet Lab’s Ancestry Analysis Certificate reports breeds in your dog’s genetic background in order of prevalence; a paragraph about each breed as well as a behavior, health and personality summary are included.
A report from DNA Print Genomics includes genotypes, ancestral population results and 15-N population results. Ancestral population results indicate your dog’s relationship to each of the four most basic branches of the canine family tree—wolf-like, hunters, herders and Mastiffs—while 15-N population results are a set of numbers that compare a sample to groups of breed signatures in the database. Each owner is given an ID and password that are used to query the online database for matches to specific breeds.
The Mars Veterinary report presents images of the breeds present in your dog’s ancestry, with the relative size of the image indicating the prevalence of each breed. An appearance, behavior and history section describes characteristics of each identified breed that might be seen in your dog.
As the companies run more tests and add breeds to their databases, the accuracy of the results may improve. It is also possible that even more companies will enter the canine breed ancestry DNA test field. When deciding where your dog’s sample will be submitted, determine what you want to learn and educate yourself about each of the tests. All of the companies agree that working with your veterinarian will ensure that you get the most out of your results. Who knows, choosing the test that best matches your needs could do more than just earn you a victory in the great German Shepherd/Doberman Pinscher debate!
*None of these tests are designed to identify purebred dogs, and the AKC will not accept test results for registration purposes.
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Dog's Life: Lifestyle
The doomed ship's survivors included three canines
April 2012 marks the 100th anniversary of the sinking of the Titanic. Knowing that only 31 percent of the ship's human passengers survived, I was surprised to learn that three dogs made it safely to New York.
Only first class passengers were allowed to bring dogs on the voyage and many belonged to prominent families. There were 12 confirmed dogs on board the Titanic including a Toy Poodle, a Fox Terrier, a French Bulldog and millionaire John Jacob Astor's Airedale named Kitty. The three survivors were all small enough to be smuggled onto the lifeboats—two Pomeranians, one named Lady, and a Pekinese named Sun Yat-Sen who belonged to the Harpers, of publishing firm Harper & Row.
Most of the dogs did not live in the cabins with their family and instead were cared for by crew members in the ship's kennel. Some of the pets were even insured, but mostly because they were considered property. However, that wasn't the case for all of the dogs aboard the Titanic.
There are many heartbreaking stories that came out of the disaster, but as a dog lover, I'll never forget the one about Ann Elizabeth Isham and her beloved Great Dane. Although many passengers regarded their animals as material possessions, Ann was said to have visited her dog every day at the ship's kennel.
Legend says that when Ann tried to evacuate with her Great Dane, she was told that he was too large. Ann refused to leave without him and got out of the lifeboat. When a recovery ship toured the wreckage days later, the crew spotted the body of a woman holding onto a large dog. It's assumed that the bodies recovered were that of Ann and her Great Dane, but the information is unverified. However, whoever the woman and dog were, one thing is for sure—they were there for each other until the very end.
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