(University of Chicago Press)
June 10 2016
To Raymond and Lorna Coppinger and their chief hagiographer at the New York Times, James Gorman, who wrote a fawning profile of the pair recently, the vast majority of the world’s 1-billion dogs all look alike because they have evolved to fill the ecological niche of village dump-diver or biological garbage disposal. Like all of Raymond Coppinger’s books, many of them co-authored, What Is a Dog? is a reductionist work of illogic that relies on simplistic scientific arguments and pre sent ism, manifest here in the assumption that the present circumstances of street dogs or village dogs have always been thus. The argument is grounded in Ray Coppinger’s belief that dogs cannot possibly have evolved from gray wolves because they look nothing like large northern wolves who feed on caribou, moose and other large animals. Were he to compare those 30-pound street dogs to the small desert wolf, he might find something different.
At the base of this book lies the Coppingers’ notion—wrong in all regards—that dogs are a species unto themselves and began to appear some 7,000 years ago, a time coincident with the first dog burials. The first dog burials in the archaeological record date to 12,000 or more years ago. The Coppingers also misrepresent or ignore evidence that dogs evolved from a gray wolf, most likely a now extinct subspecies, and continued to crossbreed for thousands of years with wolves who arose about the same time dogs did. Genes flowed from wolves to dogs and dogs to wolves. In some parts of the world, the crossbreeding continues. In the Caucasus, for example, wolves and livestock-guarding dogs are still interbreeding.
The Coppingers take what can only be described as an ahistorical view of the dog-human relationship. They seem to believe it has always resembled the current model of the dog occupying the niche of garbage disposal and occasional early warning system for incoming human or nonhuman predators. Some attention is paid to the system of transhumance—the seasonal movement of sheep between mountain and lowland pastures—but nearly nothing is said about other historic and traditional uses of dogs in particular cultures. Having spent most of their book arguing that 85 percent of the world’s 1-billion dogs are street/village dogs—the rest being human-created purebreds or their crosses—that all look the same and occupy the same niche, the Coppingers leave themselves little room for a serious discussion of just who dogs are.
News: Guest Posts
An international group of scientists proposes dual domestication from wolves.
June 4 2016
Among the many hotly debated topics related to the appearance of dogs in the lives of humans is how often and where it first occurred. In their landmark 1997 paper on dog origins, Robert K. Wayne, Carles Vilá, and their colleagues made the case for multiple origins, but many other students of dog evolution, including Peter Savolainen, a co-author on that paper, have repeatedly and strongly argued for a single place of origin.
In this week’s Science magazine (June 3, 2016) [the article is available here, gratis], Laurent Frantz of Oxford University’s ancient dog program, writing for more than a score of his colleagues from institutions around the world, presents the case for dual domestication of Paleolithic wolves in Western Eurasia and Eastern Asia. According to this hypothesis, a now extinct ancestral wolf split into at least two genetically distinct populations on opposite sides of the Eurasian continent where they encountered and joined forces with humans to become dogs.
Frantz and his coauthors pin much of their argument on analysis and comparison of the fully sequenced genome of a 4,800- year old dog unearthed at Newgrange, Ireland, to other ancient and modern dogs and modern wolves. They found it retained “a degree of ancestry” different from modern dogs or modern wolves. Using that and other evidence the researchers argue that the most comprehensive model for the appearance of the dog involves at least two domestication events 15,000 or more years ago. Frantz writes: “The eastern dog population then dispersed westward alongside humans at some point between 6,400 and 14,000 years ago, into Western Europe (10,11, 20), where they partially replaced an indigenous Paleolithic dog population. Our hypothesis reconciles previous studies that have suggested that domestic dogs originated either in East Asia (9, 19) or in Europe (7).”
I asked Greger Larson, co-director of the Oxford project and corresponding author on the paper, just what were the boundaries of “Western Eurasia,” comprised apparently of Europe and the Middle East, and “Eastern Asia?” He answered in an email that the boundaries were left deliberately vague because where wolves became dogs remains unknown, like the date itself.
In Science, Frantz writes: [W]e calculated the divergence time between two modern Russian wolves used in the study and the modern dogs to be 60,000 to 20,000 years ago.” The first number puts the dog in the time when Neanderthal was still the big kid on the European block, raising the possibility that Neanderthal had protodogs or that early modern humans came to Europe with dogs or soon allied with wolves. Either of the first two prospects must have set off alarms in some circles for Frantz cautions that those dates should not be taken as “a time frame for domestication” because the wolves they used may not have been “closely related to the population(s) that gave rise to dogs.”
Fundamentally, this paper is at once a bold attempt to come up with a workable hypothesis to explain the appearance of the dog in human affairs and a tentative step into troubled waters. Left unanswered are virtually all outstanding questions regarding the who, what, when, where, and why of the transformation of wolves to dogs. Geographically all it does is exclude Central Asia. Whether it does so wrongly may depend on how you define Central Asia geographically.
What makes it bold and radical even is the suggestion that early humans and wolves could have gotten together wherever and whenever they met on the trail of the big game they were following. There are many reasons for that including similar social and familial cultures, but humans and wolves could have joined forces to have become more successful hunters. We learn from Wolves on the Hunt: The Behavior of Wolves Hunting Prey by L. David Mech, Douglas W. Smith, and Daniel R. MacNulty (Chicago, 2016) that while wolves appear excellent at finding and trailing game, they are not very good at making the kill, succeeding perhaps half the time. It is dangerous work at which humans with their weapons excel.
Imagine the scene: Human hunters locate wolves on the hunt by watching ravens who are known to follow them. Human hunters move in for the kill and take as many animals as they can. If smart, they might share immediately with the wolves. If not, the wolves might consume what the humans do not carry off or follow them back to their encampment to take what they can.
The rest is a tale of accommodation through socialization—the ability to bond with another being—and all that entails.
This article originally appeared in Psychology Today's Dog's Best Friend, used with permission.
News: Guest Posts
December 16 2014
One of the most shared recent articles in the New York Times was one about a “wrong dog” and how the op-ed blogger felt she was wronged by agreeing to adopt a young dog from a rescue group. I was going to write about this but then our good friend, and former Bark science editor, Mark Derr, wrote a great post for Psychology Today that brought up all the points, and then some, that I had wanted to make. He kindly allowed us to cross post his article:
The New York Times ran a opinion piece on Saturday, December 13, by Erica-Lynn Huberty on the trauma caused when a well-meaning young couple bring a sweet young rescue dog into their home who turns into a cat-killing maniac. The essay, “The Wrong Dog,” serves as a sobering reminder that not all found dogs fit as seamlessly into their new homes as Arthur, the Ecuadoran stray who joined a team of Swedish adventure racers and traveled several hundred arduous kilometers with them last month. The team captain then sought and won permission to take him home to Sweden, and their story went viral.
Arthur’s story raised several questions in my mind: How frequently can dogs be said to choose their human companions, what criteria do they use, and what is their success rate? I have several friends who literally rescued dogs off the street, in one case the Brooklyn Bridge, and took them home to discover they had a friend for life.
Is it merely random chance that a dog and man or woman should meet and become instant friends? I think that both are choosing—the human to save a fellow creature in distress; the dog to find a loyal companion. Any dog dumped in the road would want that but be suspicious, too, I should think.
People I know with multiple dogs often have dogs dumped near them by neighbors who assume they will take the dog in. They do and if it doesn’t fit into their existing “pack,” they will find the dog a home. The private placements I know of have worked well—on occasion spectacularly. But dogs who go that route are the lucky exception among the abandoned millions.
The apparent ease with which human and dog share affection and respect casts light on why wolves and humans teamed up initially. Though the reasons remain mysterious, they clearly, I have long suspected, have to do with the ability of individuals from both species to form lasting bonds of friendship with someone other than their own kind and to do so voluntarily, as adults, as well as children and puppies.
Whatever mutations governing sociability occurred to make dogs, at least one must have involved fixing them as dominate in the dog genome—or so it appears.
But there are times human and dog don’t match up well, and unless something is done, the results can be tragic. Many of the failures in that relationship seem to arise from a lack of forethought on the part of the human, a fundamental failure to think through and find ways to meet the animal’s need for exercise, social contacts with people and dogs, consistent treatment and mental stimulation.
The central problem with Huberty’s essay lies in her argument that nothing short of ditching the dog when she first started acting oddly would have prevented the catastrophe that occurred. They would have done that had they known that some dogs are unfit for adoption, and no amount of training, discipline, or coddling will change that.
“We let ourselves believe that beneath our rescued puppy’s strange, erratic behavior was a good, loving pet,” Huberty writes. The truth was the opposite.
The back story is common enough. Having become smitten with a five-month old Lab mix, Huberty and her husband, decide to have her share their home with their three cats, a female dog, and two children.
From her arrival, the new dog, Nina, showed a defensive/possessive aggression that led Huberty to seek more information from the group who rescued her.
Huberty says that she and her husband followed the advice of Cesar Millan, “the Dog Whisperer” to create a “loving but disciplined environment.” Nina responded by attacking a cat and biting Huberty when she intervened.
In response, Huberty called the woman who gave them Nina. She agreed to pay for a trainer, who proved to be the anti-Millan. She advocated a rewards-based approach rather than “discipline.” The essay takes an odd turn here as Huberty calls the rewards-based method ‘coddling” while appearing to indicate that it was working up to a point.
Nina would go along being a normal, playful puppy. But at times, out of nowhere it seemed, she would snap at me or Alex and, once, at our son,” Huberty says, “She would suddenly cower and growl. It was like a switch flipped, yet we couldn’t figure out what had done it.”
Nor do they try to find out. Dogs do not usually change their behavior that rapidly and dramatically without reason. That could very well be an underlying pathology that a thorough examination by a veterinarian might reveal. Indeed, Huberty gives no indication that she ever took the dog to a veterinarian—the first stop a new dog or cat companion should make.
If no physical reason for the behavior can be found, the next stop is to consult a board-certified veterinary behaviorist. There are not many in the country but your veterinarian should help arrange a consultation.
Huberty blames the dog, the woman who gave her the dog, the trainer—everyone but herself and her husband—and Nina herself for her failure to fit seamlessly into Huberty’s home. From this experience, she draws the conclusion that some dogs are just unsuitable for living with humans. That might be the case but there is no proof of it here.
Maybe we should seek ways to allow more dogs to choose their human companions. I have a notion they would do a better job of it. “And when they don’t fit in they may be saying ‘wrong family,’” said my fellow Psychology Today blogger Marc Bekoff after reading “The Wrong Dog.” “Living with a dog is a two-way street and assigning unilateral blame gets us nowhere and once again leaves the dog out in the cold. This sort of ‘musical dogs’ is bad for the dog, as much research and common sense tell us.”
Nina might pay with her life for human miscalculations and failure to seek professional help.
News: Guest Posts
On the whole, human breeders have not improved on nature.
May 16 2014
On a flight last year, I sat next to a woman from India on her way to London from New York, where she had been visiting her first grandchild When she heard I wrote about dogs, she turned her attention to the one aspect of her daughter and son-in-law’s life she could not understand—their dog. On her walks around Central Park with the dog and her granddaughter, the dog drew the most attention and comment.
He was a black, blue-eyed French Bulldog for whom they had paid $3,000 to a veterinarian/breeder in New Jersey. They had already spent that much again on veterinary bills in less than a year.
Defenders of the purebred dog industry talk a lot about responsible breeders, and I once tended to follow their lead. But shortly after the Atlantic Monthly published my article, “The Politics of Dogs,” in March 1990 [It is hard to find on-line because of a class-action lawsuit regarding electronic rights, but it can be found.], I began to hear from people who had, like this couple, gone to a responsible breeder only to end up with a dog with problems. Many of the genetic conditions to which pedigreed dogs are prey do not follow strict lines of inheritance—they skip generations or move through aunts and uncles. They do sort by breed, but that is because those breeds have arisen from a small number of founders—in short, they are inbred, dangerously so, and this tight knit extended family shares most strongly genetic diseases and physical characteristics. Outer beauty conceals inner flaws.
French Bulldogs, for example, are prone to von Willebrand’s Disease, a blood disorder, as well as spinal problems, a cleft palate, and heat stroke. Many airlines no longer carry brachycephalic breeds—those with pushed in faces—because they have a tendency to die in flight from breathing problems related to overexcitement.
In light of that, the woman asked, why do so many people spend so much money on these dogs? This conversation occurred months before the sale of a Tibetan mastiff puppy at a luxury pets’ mart in Hangzhou, China, to a Qingdao property developer for 12-million yuan (about $1.9 million), reportedly a record price for a dog. It is easy here to invoke the 19th and early 20th century economist, Thorstein Veblen and his theory of conspicuous consumption.
To Veblen, such dogs are objects of conspicuous consumption, animals with no intrinsic value that nonetheless are made valuable by the fact that someone goes to great lengths to obtain and maintain them despite or because of the expense involved in doing so. Put another way, possession of such a being marks you as a person with so much money that you can obtain and maintain an animal with no useful talent.
That would certainly be the case with the Tibetan mastiff, which according to some assays is merely a reconstruction of a once mighty landrace of large livestock protection dog, which it resembles the way a teddy bear resembles a grizzly cub.
Clearly spending that much on a dog must be considered conspicuous consumption of the most extreme sort. It is also a mordant commentary on the Chinese Revolution, for half a century ago, Mao Zedong sought to rid China of pet dogs he considered objects of bourgeois recidivism—that is, conspicuous consumption.
Currently the recidivists have won. In China and other countries with a growing urban middle class, people are buying more and more dogs, eschewing their local dogs for Western pure breeds. To them, the pedigree signifies quality.
When Veblen used the Pekinese as an example of an object of conspicuous consumption, purebred dogs were relatively new on the scene and well beyond the means of most people. A century later, the dogs are no longer rare, nor are their prices, even at $1,000, so outrageous, especially when buyers are convinced they are getting excellent bloodlines, superior quality, and specific behavioral characteristics.
Those beliefs fuel demands for purebred dogs produced by commercial breeders—let’s just call anyone engaged in the large scale “production” of puppies for profit, a commercial, or mass, breeder, and recognize that some are better than others, which is not an endorsement of any of them.
Demand for purebred dogs shot up following World War II when returning veterans, establishing their lives in burgeoning suburbs, sought them out as accompaniments to their new homes, cars, and families. The pedigree provided by the rapidly expanding American Kennel Club, the largest registry of dogs in the world, proved these acquisitions were not the old family mutt, but refined and sophisticated pets. Demand fueled the growth of mass breeders, pet stores, dog shows, regulations to fence and leash dogs, and unwanted dogs.
I estimate that by some point in the 1990s half of all dogs In America were purebred, and a great many of them were from mass breeders. That was a problem because they too frequently bred dogs without regard for their temperament or genetic soundness and failed to socialize them during the critical first three to four months. If dogs are not socialized to humans during that time, they might have difficulty ever becoming fully socialized and often have behavioral issues.
The problem with these breeders has been known for decades and several national animal advocacy groups have campaigned against them for years without much result. Although there are many political explanations for the failure to end the retail trade in dogs, these groups have not invested in the sort of intense, dedicated campaign required to shut it down.
Instead, we get things like the Humane Society of the United States forming a group, Breeder’ Advisory and Resource Council, to advise it on matters relating to responsible dog breeding.
Mass breeders are a significant part of the problem of purebred dogs, but not the only one. My colleague and fellow Psychology Today blogger, Marc Bekoff, has argued for a full halt to breeding more dogs as long as millions of perfectly fine, adoptable animals await new homes in shelters or the homes of breed rescue group volunteers. With so many dogs in need, he says, the compassionate and humane thing for someone wanting a dog is to adopt one.
I believe he is right, and I would add that the breeding of dogs as it is now done should stop until ways can be found to minimize the risk of a dog being born with an inherited deformity or illness. That includes behavioral problems. These conditions disproportionately seem found in puppy mill dogs, but not exclusively. Some of these conditions, especially cytoskeletal ones like brachycephaly are so extreme that puppies must be delivered by caesarean section and subsequently have difficulty breathing normally and dissipating heat. The book on these inherited ailments is long and growing longer. Most are due to the heavy inbreeding and common use of favored sires in breed formation.
It is worth remembering here, that breeds are formed through consolidation from an existing population, when a few animals are used to create the Platonic ideal of the ‘breed,’ and through amalgamation, in which representatives from several similar landraces are crossed to create the perfect representative of all.
No matter which method was followed, the resulting dog was said to represent the breed in its pure essence and be more intelligent and talented than any of its naturally breeding predecessors. With few exceptions involving specialized behaviors that have been enhanced through selective breeding, that is untrue. Nonetheless, like mantras, the histories of these new breeds and accounts of their prowess were repeated so often, they became truth—to everyone but the rulers of the American Kennel Club. For decades they have publicly maintained that the AKC issued pedigree proving that for three or more generations the dog in question is an official Chesapeake Bay Retriever or whatever the case may be does not represent quality, does not guarantee that the dog is healthy or possessed of a good temperament. They did that because they wished to avoid possible consumer lawsuits involving dogs with serious defects and flaws.
Despite those disclaimers, the AKC has continued to a promote the virtues of purebred dogs, like the problematic French Bulldog, the eleventh most popular dog it registered in 2013, and the larger English bulldog. It was the fifth most popular breed registered in 2013, even though nearly 72 percent of the bulldogs evaluated by the Orthopedic Foundation for Animals had crippling hip dysplasia; none could whelp naturally.
The argument is sometimes made that breeds are important biological artifacts but, in fact, over the decades breeders have altered their appearance—and perhaps their behavior—and perhaps their behavior—substantially. Many breeders say that they are attempting to improve the breed they love, but the very notion that a breed needs improvement suggests that it has problems.
Few if any breeders can predict that all puppies in a litter will be free of congenital defect, but in many cases the odds are stacked against them from the start by a plethora of problems and conditions associated with their breed. Breeders and kennel clubs should focus on ridding the breeds they love and promote of those inheritable conditions, and the way to do that is to stop engaging in dangerous breeding practices and to avoid breeding dogs who have them in their bloodlines.
Dogs deserve no less.
This post first appeared on Mark Derr's blog, Dog's Best Friend on Psychology Today. Used with permission.
News: Guest Posts
“Do As I Do” scores high
March 8 2014
A rambunctious five-year-old Labrador Retriever who until a few months ago knew not a word of any language, obeyed no command, charged around the house or zipped through any hole in the fence before one could utter the name he didn’t seem to recognize has become my 91-year-old mother’s great and constant companion. He sits or lies by her when she is sitting or lying down. He moves with her when she goes somewhere with her walker and when she tells him to give her clear passage. He accompanies her when she walks around the pool for exercise. She says, “He is a good boy.” My mother has never trained a dog. She had a nice trained dog once, but she had been trained by someone else and given to her.
But Rocky, as he was named by my mother’s granddaughter, received no formal instruction from any source. He was neutered, which helped slow him down, but more profoundly, he and she opted for companionship and accommodation over ignoring each other. She talks to him constantly, telling him what she wants him to do. If she praises him, she is not effusive. She may occasionally slip him some food when she is cooking, and he will if given a chance steal her breakfast bagel. There is no system to it, but there is consistency.Top of Form
More than a few dog trainers who follow behaviorist principles that require a stimulus, a reward or punishment, for learning to occur would argue that Rocky is untrained—that is that he still will not perform on command the actions demanded of him—except he comes when called. He moves when told. He tells my mother when someone is at the door and stands by her when she opens it, thereby providing at least the illusion of protection. If that is not training, what is it?
My friend and fellow Psychology Today blogger, Marc Bekoff (“Animal Emotions”), might call the process dog teaching or dog learning.
It might not be as quick or as systematic as one of the common schools of training, including those that use electric collars and choke chains and those that rely on clickers and food rewards or other positive re-enforcers. But then again the results might be quicker, deeper, and longer lasting.
I have seen no statistics on the numbers of dogs educated in this fashion, but I imagine it is substantial. Essentially it relies on the dog’s innate curiosity, desire to please, and recognized ability to imitate behavior and recognize words and emotions, traits which arguably thousands of years of living with humans have served to enhance. It also requires the human have an interest in being with the dog and interacting with him or her in a meaningful way—what used to be referred to as “quality time” with the hound. Praise and rewards are meted out more according to the person’s nature than any program or schedule. They do not have to involve food. Our Kelpie Katie was unmotivated by food—she would ignore food rewards—but when a tennis ball appeared she went on high alert. Even then the ball was not essential to her learning something.
This intuitive style of dog teaching is not without its intellectual underpinnings thanks initially to Edward Tolman in the first half of the last century. He proposed that learning had intrinsic value and that people and animals could learn in the absence of immediate rewards—latent learning it is called. That idea underpins what is called the social theory of learning, which also views learning as a social endeavor that can involve imitation of behavior that is demonstrated or verbally described.
In an article in the January 28, issue of Applied Animal (Behaviour Science, entitled “Should old dog trainers learn new tricks? The efficiency of the “Do as I do” method and the shaping/clicker training method to train dogs,” Claudia Fugazza and Ádám Miklósi of the Eötvös Loránd University in Budapest, Hungary, look at a canine system of social learning that relies on the dog’s great capacity for imitation called Do As I Do (DAID) compared with clicker training, which relies on the timely delivery of rewards to employ the dog’s associative abilities in shaping its behavior. (The article is only available by subscription, but here is the Abstract.) The clicker becomes a stand-in (secondary re-enforcer) for the actual re-enforcer, usually food. Clicker training is individualized instruction that requires the dog to figure out what earns rewards.
Fugazza, a graduate student in ethology developed Do As I Do in order to study social learning in dogs. To do that she had to develop protocols for teaching them. Judging from its success, it should gain a wide following. In this method, trainers, usually the dog’s primary human companion, use standard reward-based techniques to teach the dog to associate a small number of gestures with the command, “Do It!” The dog is then shown a new task and taught to perform it upon being given that command.
For this study, Fugazza and Miklósi compared the speed in learning three sets of tasks of increasing complexity, from knocking over a glass (simple) to opening or closing a locker or drawer (complex task) to a sequence of actions, like hopping on a chair and ringing a bell or opening a locker and removing a purse (compound). Objects were involved in each task that were not considered part of the family dog’s normal repertoire so that mastery of the task could be construed as learning. In the simple task there was no difference in performance between clicker-trained dogs and Do As I Do dogs, but that changed as the tasks became more difficult. Do As I Do dogs performed noticeably better, with more of them learning the task in the allotted fifteen minutes than clicker-trained dogs.
No one knows how the dogs are making the connections, and in their conclusion Fugazza and Miklósi thought it more important to downplay that result in favor, Miklósi said in an email, of providing trainers with as many methods as possible so they can choose the one best suited to their needs.
That is a tactical decision rather than a scientific one. It is grounded in the recognition that, especially commercial dog trainers and trainers of working and service dogs, like to use what has worked for them in the past with the kind of dog on which it has worked. That is one reason punishment-based forms of dog training persist.
For home schooling, time, patience, devotion—and a daily reminder of who has the big brain—are the keys to success and those come from discipline we often need more than the dog.
Used with permission of Mark Derr and Psychology Today, see more from Mark Derr’s blog “Dog’s Best Friend.”
Also see http://thebark.com/content/dogs-are-asked-just-do-it
News: Guest Posts
A fine memoir of a road trip with dogs to the World Sheepdog Trials
June 23 2013
Not far into Mr. and Mrs. Dog, Donald McCaig says of himself and his talented “Blockhead” of a Border Collie, Luke, the male of the title: “I’ve never done as well with Luke as a better handler might have, but Luke adores me. When I go out at 2 a.m. to check lambing ewes, Luke comes too. When I wake with the night sweats, Luke wakes. He thinks I am a better man than I am. If I sold him, his earnest doggy heart would break.”
It is a tribute to McCaig’s capacity for self-reflection and humor that he is willing to admit his own failures as an occasionally over anxious sheepdog handler. He knows that dogs are not machines and we are not infallible. Ultimately all you can do is the best you can do under sometimes disastrous circumstances.
Upon reaching 68 years of age half a decade ago and finding himself with two quality border collies in their prime, McCaig decided the time had come to launch a campaign to fulfill his dream of the worlds.
Traveling 34,000 miles in his twenty-year-old car, McCaig, Luke, and June (Mrs. Dog) compete in sheepdog trials around the country hoping to compile enough points to secure invitations to join the American team in Wales. At the last minute, June garners the invitation, and Luke gets to compete as McCaig’s second dog.
If his best-selling Nop’s Trials is McCaig’s contribution to “lost dog” literature—think of Jack London’s Call of the Wild and White Fang—Mr. and Mrs. Dog: Our Travels, Trials, Adventures, and Epiphanies is his homage to an equally venerable tradition, the “the dog road trip,” of which John Steinbeck’s Travels with Charley is perhaps most famous. McCaig is on the road not only to qualify for the worlds but also to broaden his dogs’ experience of different sheep and environments—in a fundamental sense to educate them so they will be better able to cope with situations and varieties of sheep they have not seen before.
Although June pulled them through on cumulative points for the year, her most memorable performance came at a trial in West Texas when she decided to forego herding sheep and goats in favor of far bigger game--a huge, ground-thumping oil exploration seismograph truck. “June wanted, nay NEEDED to fetch that big thumping, flickering weirdness,” McCaig writes, “and nothing I said—neither my shouts nor redirects—swayed June from her goal.”
Once abreast of the thumper, June realized she had not a clue what to do with it and returned to McCaig, but there were no longer any goats to fetch. Her assault on the seismograph thumper had disqualified her.
Hoping to further his own education, McCaig periodically detours from the sheepdog circuit to visit trainers known for their skill in training methods they have developed or adapted. Along the way, he correctly points out that the battle between practitioners of what we might call punishment-based training and those who prefer awards-and rewards-directed training is now more than 100 years old.
For much of that time it appears that punishment has ruled—aversive training, as it were. McCaig himself is something of a follower of William Koehler, the Disney animal trainer from the mid-twentieth century, who developed a method of obedience training relying on long lines and various chain collars and leashes. Even today, most people attending obedience classes probably follow some version of Koehler’s method.
McCaig is looking for training epiphanies; bright moments of understanding or enlightenment that will help him better train and manage his dogs. He meets animal behaviorist Nicholas Dodman, and attends sessions given by trainers using the dog’s ‘innate’ drives, rewards, the Koehler method, and shock collars, which so engage him that he adopts the industry’s terminology and calls them e-collars.
Over the years, McCaig and I have agreed to disagree about shock collars, and in future postings, I hope to examine different approaches to training. For now, I’ll just say that people searching for a blanket endorsement of shock collars or other training devices or methods will not find them here—with the possible exception of the thirty-foot long line, which need not deployed in punitive fashion.
McCaig’s book arrived shortly after I had visited my favorite trainer, Lourdes Edlin. She is one of those gifted people who will have a dog literally eating out of her hand within minutes of meeting it. She understands that to train a dog, she must learn what motivates it—food treats in many cases, but in others a ball or Kong® or simply praise.
Edlin said that she was growing tired of teaching people basic obedience—sit, stay, heel, come—and becoming more focused on “teaching people how to do things with their dogs.” The basics would follow from that.
I was reminded of Edlin’s comments when I read McCaig’s reflections on his forays into the world of training. “Though each trainer believes his or her method is best, I don’t think it matters which method the pet owner adopts so long as that owner finds a capable mentor and sticks with the training,” he writes. ”Eventually you will learn to see your dog and when that happens the richness of your and your dog’s lives will tell you what to do next.
“Neither Luke nor June was ever trained to ‘heel’ nor ‘sit’ nor ‘stand for examination.’ They have never retrieved a ball or dumbbell. They rarely play with each other and never play with other dogs. Yet they would be mannerly in any human environment. Not because they were ‘trained’ for good manners, but because they were properly socialized, exercised daily, and have a job—stock work. Mannerliness is a by product of that training.”
A few paragraphs later, he concludes, “Have the highest expectations, do the work, and your dog can walk at your side anywhere on earth. He’ll become the dog you’ve empowered to change your life. As Luke and June have changed mine.”
McCaig’s account of the trio’s trip to Wales is informative, amusing, and somewhat sad. The two males manage to win a local Welsh competition, the South Wales Sheepdog Trials Hafod Bridge, where McCaig penned his sheep brandishing his Stetson® hat instead if the traditional shepherd’s crook. A revolution was doubtless averted when McCaig confessed that he simply had deemed his crook too difficult to manage on the flight across the pond and he had neglected to obtain one. Clearly a telescoping shepherd’s crook is in order.
Luke, June, and McCaig washed out in the first round of the big show. McCaig blames himself for failing to meet his expectations, but he should not.
He’s written a fine book and made a most excellent life with Mr. and Mrs. Dog. Moreover, they have had many an excellent adventure. What more could a dog or human want?
This blog originally appeared on Psychology Today. Reposted with permission.
News: Guest Posts
May 22 2013
The more I consider the continuing debate over the “time” and “place” for the transformation of wolf into dog, the more I become convinced that the puzzle remains unsolved because of human devotion to a simplistic, clever-sounding idea that never made sense in the first place. As first put forth by Raymond Coppinger, that idea was that wolves feeding on the garbage piles of quasi-permanent Mesolithic villages grew tamer over the course of generations until they no longer feared or threatened humans. In the process of taming themselves, those wolves also became less fearful of and aggressive toward humans. They were cute, too. For reasons that were never clear to me, people took these cute obsequious dump divers into their homes, where they blossomed into dogs.
Coppinger pinned his argument on Dmitry Belyaev’s experiment, begun in 1959, at a Siberian fur farm, in which a group of foxes was bred for tameness alone and within ten generations was producing foxes that resembled dogs with floppy ears, piebald coats, and a high need for attention. They were juvenilized in behavior as well as appearance.
There are a number of reasons why the foxes are not a good model for origins of the dog, and I have elsewhere addressed them in detail. For now, suffice it to say that dogs arose not in quasi-permanent Mesolithic villages but in Paleolithic hunting camps. They were not sought nor selected because they solicited attention and showed no aggression—these are hardly traits of a good guard, which was one of the tasks of early dogs. Guarding remains a major reason why people keep dogs.
But the greatest problem with the self-domesticating theory is that it shuts the most creative creature on the planet out of the process. To put it bluntly, that makes no sense. Humans have always collected, tamed, and trained animals. It is inconceivable that they would ignore one as intelligent and inquisitive as the wolf.
Genomics and its offspring have shown that living organisms are not biological machines but energetic systems supported by layers of complexification. Genomics has also contributed to a more dynamic view of “domestication” as a process involving the interplay of biological, environmental, and cultural forces. The hard line between “domestic” and “wild” –always imaginary but not less real for that—has for the dog become increasingly difficult to find despite the distortions that define the current period of breedism. I am using “breedism” to refer to all aspects of the cult of the purebred dog that began to take hold about 200 years ago. Of course, there are significant differences between dogs and wolves, when they are in their own environments, but what happens when the dog goes native or the wolf becomes a lay-about?
It sometimes appears that every new find simply raises new questions while leaving old ones unresolved. That trend is apparent in two new papers by Ya-ping Zhang, a leading Chinese geneticist, who collaborated with geneticists from China and two different labs in Sweden and California on two new papers promoting Chinese indigenous dogs—native or village dogs—as the closest dogs to the ancient type.
Working with Peter Savolainen, of Sweden’s Royal Institute of Technology, Zhang has over the past decade or so insisted that dogs originated in southeast China no earlier than 16,000 years ago, and many researchers elsewhere adopted his argument despite the notable absence of dog or wolf remains from that region at that date and the presence of dog remains from other places considerably earlier.
The researchers redid the numbers using new chips that spot changes in the genome including so-called single nucleotide polymorphisms, or SNPs, and indels—insertions or deletions of small amounts of genetic material. SNPs and indels can be used to measure degrees or years of separation of discrete groups of organisms. Crunching the numbers and running regression analyses, they found that southeast China village dogs separated from wolves 32,000 years ago. There were a lot of them by then, too, they reported in an article in Nature Communications [subscription required], with Guo-dong Wang and Weiwei Zhai as first authors and Zhang as senior author—8,500 dogs by their estimate.
The new date fits nicely with some “early dogs” identified from the Altai Mountains, Belgium and the Czech Republic, although Zhang and his colleagues are not quite willing to admit that those animals are dogs. In fact, they appear to want to deal with the early date by using it to mark the beginning of a long period of self-domestication for a group of scavenging protodogs.
Zhang’s group declares: “Early wolves might have been domesticated as scavengers that were attracted to live and hunt commensally with humans. With successive adaptive changes, these scavengers became progressively more prone to human custody. In light of this view, the domestication process might have been a continuous dynamic process, where dogs with extensive human contact were derived from these scavengers much latter [sic] when humans began to adopt an agricultural life style.”
The operative words here are “commensally” and “scavenger.” Together, they say that wolves were drawn to human garbage or some other waste and so started hanging around and hunting with them but without having a discernable effect or bringing them any benefit—thus, the term “commensally”—until the biped started farming. Then the scavengers showed their true worth as crossover omnivores and became dogs.
That is not complex, but it is convoluted. At a basic level, it is not clear why protodogs could not have arrived in southeastern China from the Altai Mountain region, for example, where the people who would come to enter the New World and spread through much of the Old World as the glaciers began to retreat, had gathered, presumably with dogs some 35,000 years ago. A population of dogs and people could easily have gotten to southeast China and radiated outward from there. The much trumpeted diversity of dogs in the region could be a result not of their origins there but an accident of geography and history, including intensive breeding of dogs for food and a settlement pattern that featured many small riverine villages along the Yangtze River, one of the world’s largest.
Zhang’s defense for the lack of wolves in southeast China is that wolf populations have changed everywhere, and so no one has an ancestral wolf for study and comparison. But the Chinese indigenous dogs and a couple of related breeds, are the dogs closest genetically to wolves, and that makes them all the more important as living artifacts, Zhang and his team reason. Specifically, they looked for genetic loci that might show positive selection pressure in dogs and humans and therefor might represent parallel evolution in the two species. The genes they identified as likely candidates are involved in diet, specifically the ability to digest grains; metabolism; cancer and neurological processes, especially some involving the neurotransmitter, serotonin.
Zhang is also corresponding author with Dong-Dong Wu, both of the Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China, on a Molecular Biology and Evolution paper involving the laboratory of Robert K. Wayne, dean of canid evolutionary biologists, and several of the next generation of dog geneticists who have already published important work—Bridgett vonHoldt and Adam Boyko. They were especially focused on the prefrontal cortex and on parts of the brain that appear involved in fear response and sociability.
I say “sociability”, but, following Zhang’s lead, the researchers on these pages say “tameness,” while continuing to cling to the Soviet fox experiment as evidence that the dog was self domesticating, becoming obsequious and ingratiating and nonaggressive while eating garbage and offal. Standard descriptions of this work are abundant, and I won’t repeat them here. But it is fair to say that grand pronouncements about the working of the brain must be treated cautiously.
Most of these searches for genes involved in the transformation of wolves to dogs are based on at least two significant, faulty assumptions about the behavior of dogs and wolves. The first faulty assumption Is that wolves are now, and were in the late Pleistocene, aggressive competitors with humans. There is evidence documenting not only friendly but also mutually beneficial relationships of humans and wolves going back thousands of years. There are suggestive associations of wolf and Homo erectus remains going back hundreds of thousands of years.
The second faulty assumption is that a group of wolves effectively said to humans, “Because we like your leavings so much, we will stop vying with you and aggressing against you. We will be abject before you if you will give us excretia to eat because we cannot live by ourselves.” The question I always ask is, would you want such a creature in your house, in your bed? That is unlikely. This assumption is faulty because there is no evidence that wolves generically dislike or even fear humans. The global wolf recovery with wolves living in ever closer proximity to humans proves that wrong. It is humans who hate wolves.
That wolves and humans, similar as they are in so many ways, should make common cause, should surprise no one. Hunters study hunters. Species cooperate. It would be more aberrant if they did not. Coral groupers, Napoleon wrasse, and moray eels were recently shown to hunt cooperatively, for example. Around the world, hunting cultures had dogs that often interbred with, sometimes were indistinguishable from wolves. In the New World and elsewhere that situation was contemporaneous with the rise of multiple refined breeds in the Anglo-English speaking world.
In a real sense, then, what we call domestication of the wolf was really a rolling and flexible bringing into human culture of wolves who had the psychological and emotional capacity for sociability, for forming strong bonds not just with another individual but also with another species.
Some years ago Adam Miklosi and his colleagues compared hand reared wolves to dogs. The lengthening of the first critical socialization period and a greatly increased capacity to form strong bonds to another species were clearly central to the appearance of the dog, they concluded.
Yet for all of their problems, these two new studies are useful for their focus on indigenous dogs, the landrace dogs who although they might have several uses are generally not bred by humans to any purpose, but who still live, reproduce, and die in human society. How ancient or basic these dogs are is not really known. But they are found around the world, and I think that comparative studies of them and resident wolves and truly self-sustaining feral dogs, where they still exist, will prove most interesting. The same applies to comparison of DNA from ancient dogs and wolves. We do not yet see them clearly.
This article first appeared on Dog’s Best Friend at Psychology Today, it is used with persmission.
Mark Derr is the author of six books, ranging from Some Kind of Paradise, an environmental history of Florida, How the Dog Became the Dog, Dog's Best Friend, and A Dog's History of America. As an expert on the subject of dogs, he has been a guest on such programs as The Charlie Rose Show and Fresh Air. His articles and opinion pieces have appeared in the Atlantic Monthly, Wall Street Journal, Huffinton Post, Natural History, The Bark, Smithsonian and The New York Times. He lives in Miami Beach, Florida.
News: Guest Posts
March 8 2013
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
Celebrating the bicentennial of the father of evolution
February 27 2009
Early in The Descent of Man, and Selection in Relation to Sex, Charles Darwin uncorks a passage to illustrate the capacity of dogs to love that is guaranteed to break the heart of all but the most unfeeling cad, and one that should hang over the door of every laboratory engaged in experiments with animals. “In the agony of death a dog has been known to caress his master,” he says, “and everyone has heard of the dog suffering under vivisection who licked the hand of the operator; this man, unless the operation was fully justified by an increase of our knowledge, or unless he had a heart of stone, must have felt remorse to the last hour of his life.”
A number of dedicated readers objected when those words and a slightly different version of this essay ran in the Winter 1999 issue of Bark, saying that the editor and I were forcing them to look at the human capacity for cruelty to dogs and other animals with which they were all too familiar. Darwin’s words remain because they bespeak much about the man whose bicentennial birthday we celebrate this year—his clear eye, his intelligence, his compassion, his deep respect and love for nature and its fruits, and his conviction that we are related fundamentally to all life. Sadly, that Darwin is nearly unknown in an America where a significant number of people believe Darwin’s theory of evolution by means of natural selection compares unfavorably with a distortion of religion and science called creationism, or intelligent design.
A casual reader of Darwin’s collected works might assume that the man was obsessed with pigeons, which he avidly bred and studied as models for his theories. But closer examination reveals the full depth of Darwin’s affection for all animals and his special love for dogs, who bound through the pages of his work, illustrating his theories of evolution and domestication as well as animal intelligence and emotion. After disembarking the Beagle in 1832 (although purely coincidental, what better name for a ship on which a naturalist is sniffing out the mysteries of evolution) to explore Patagonia and environs, he made a special trip to investigate the shepherd dogs of South America, described a few years earlier by the great French naturalist Alcide Dessalines d’Orbigny.
I suspect that Darwin, a proper skeptic, wanted to determine for himself the veracity of d’Orbigny’s account of these mongrels: Suckled on ewes’ milk and raised with sheep from infancy, they traveled with their flock, taking it to pasture, bringing it home, protecting it from marauding wildlife and dogs. In their spare time, they hunted partridges and jaguars for their masters, who abused them horribly, even to the point of slashing them with knives.
In The Voyage of the Beagle, Darwin doesn’t confirm the hunting and slashing, but he does observe that, after the shepherd dog brings the flock in, he goes to the house for a piece of meat and then skulks away with it “as if ashamed of himself,” pursued by “tyrannical” house dogs. But upon reaching his flock, the dog, who, like all his kind, was castrated at an early age, turns and routs the house dogs with a bark. Darwin saw the whole performance as an example of the dog’s powers of affection and association, what today we call bonding and behavior modification by castration. It is a lesson everyone who considers the big white guarding dogs genetically preprogrammed to protect sheep should consider. (Or they can go look at Navajo sheep dogs, raised in this old Spanish way.)
Unfortunately, Darwin, like those other intellectual giants Marx and Freud, is seldom read but frequently invoked— wrongly—by his proponents and opponents alike. His classic phrase, “natural selection,” means simply, as he explained in The Variation of Plants and Animals Under Domestication (Vol. I), that those individuals “best fitted for the complex, and in the course of ages changing conditions to which they are exposed, generally survive and procreate their kind.” Those who survive to pass their genes on to the next generation are by definition the fittest. They need not be the biggest, strongest, boldest, brightest, baddest or best at corralling millions in the stock market. Indeed, there are circumstances in which the meek and cautious survive while the marauding bullies perish, victims of their own aggression.
Darwin believed in the power of dogs to help shape human evolution, writing in The Descent of Man: “The strongest and most vigorous men—those who could best defend and hunt for their families, who were provided with the best weapons and possessed the most property, such as a large number of dogs or other animals—would succeed in rearing a greater average number of offspring than the weaker and poorer members of the same tribes. There can, also, be no doubt that such men would generally be able to select the more attractive women.”
Operating slowly over a long time frame, natural selection is more potent than what we humans can accomplish through selective breeding of domestic animals. Breeders can’t create new traits for domesticated species, Darwin said, they can only concentrate or de-emphasize those already present. Through “methodical,” or conscious, selection, a breeder decides what characteristics he wants and then strives to reach them. “Unconscious” selection, on the other hand, involves “the preservation by man of the most valued, and the destruction of the least valued individuals, without any conscious intention on his part of altering the breed.”
Usually the two worked in tandem in subtle ways. By Darwin’s time, several Greyhound breeders had crossed their dogs with Bulldogs to add courage, Pointers were crossed with Foxhounds for speed and agility (to match faster horses), and Bulldogs were downsized after the demise of bull baiting. The breeders doubtless were seeking to match the fastest dogs, for example, but they could hardly foresee what the animal would look like, and it was, Darwin says, different from the old hound.
Absent any knowledge of genetics, Darwin demolishes the practice of inbreeding (called “interbreeding” or “breeding in to in”), saying it arises from a misbegotten belief in “purity of the blood,” and extols the virtue of crossing and hybrids. Inbreeding may appear to work, he says, but in the long run it produces weak, infertile animals, prone to malformation and disease. On the other hand, too much outcrossing leads to reversion to the basic dog. The key to success was to choose the best, the fittest, and proceed. In The Descent of Man, he asserts that “hardly any one is so ignorant as to allow his worst animals to breed,” meaning those who were unsound mentally or physically. (More than a few modern breeders should take heed.)
Although Darwin frequently used dogs as examples, they presented difficulties not found in pigeons. He recognized that variability was essential to the adaptability of wild species, and that domestication, for reasons we still don’t understand, unloosed the variability inherent in the wild progenitor of an animal. But the astounding variety of dogs, from the noble Greyhound, “the perfect image of grace, symmetry, and vigor,” to mutant Bulldogs, with their exaggerated heads and undershot jaws, and the turnspit dogs, with their dwarfed legs (they were literally used to turn spits), befuddled him. He could only conclude that they must have come from several different wild species, primarily the wolf and jackal, a notion we now know is incorrect.
With varying degrees of success, humans have attempted to reorder the behavior of those wild progenitors. In On the Origin of Species, Darwin observes that pointing, encircling a herd or flock, and retrieving are among the general wolfish hunting and puppy-rearing traits that have been “rigorously” selected for by nature. “Less rigorously,” or less precisely, humans had concentrated one or more of those traits in specialized breeds of dog for herding, pointing and retrieving. Other changes were more profound. He writes in The Descent of Man that dogs “may not have gained in cunning, and may have lost in wariness and suspicion, yet they have progressed in certain moral qualities, such as in affection, trust, worthiness, temper, and probably in general intelligence.” They had also become more biddable, or trainable.
Underlying much of Darwin’s thought was his profound belief, shared with other 18th- and 19th-century naturalists, that animals differed from humans largely in degree, not kind. He argued that as one moved from lower to higher orders of animals, the influence of instinct declined and intelligence increased, and dogs and horses were near the top of that hierarchy. In fact, Darwin believed domestic animals were generally more intelligent than their wild progenitors, a notion that in this century has been reversed—to the detriment of all animals.
In the opening chapters of The Descent of Man, he brilliantly argues, using dogs as prime examples, that animals feel “pleasure and pain, happiness and memory.” They inherit the capacity for terror, suspicion, fear, deceit, timidity, bad and good temperament, rage, and vengefulness. More significant, they possess the powers of reason, imagination, love, jealousy and pride. They believe in the supernatural: “There must be something special, which causes dogs to howl in the night, and especially during moonlight, in that remarkable and melancholy manner called baying.” They are also religious in a way, substituting the master for God. Not possessed of human language and learning, they nonetheless communicate—who among us can’t recognize the meaning of our dog’s barks, chortles, growls, bays, yodels and howls?—and learn in their world. When in response to a whispered, “Where is it?” a dog charges from to tree to tree, she proves that she has a notion that there is something to hunt or fetch; thus, she engages in a form of abstract thought.
Dogs, and other animals, possess a sense of beauty, or aesthetic appreciation, although mostly confined to sexual attraction. Their moral sensibility is manifest in their knowledge of right and wrong and their assistance to their family, pack or herd. But they are also altruistic: “It must be called sympathy that leads a courageous dog to fly at any one who strikes his master, as he certainly will.” In another instance, Darwin tells of a story Audubon related —“that he had reared and tamed a wild turkey who always ran away from any strange creature. This bird ultimately escaped into the woods. Some days later, Audubon saw what he thought was a wild turkey and ordered his dog to give chase. But the bird did not run away, and the dog, when he came up, did not attack the bird, for they mutually recognized each other as old friends.”
Darwin admits that animals may lack self-consciousness, the ability to reflect on the meaning of life and death, their place in the cosmos. “But,” he then slyly asks, “how can we feel sure that an old dog with an excellent memory and some power of imagination, as shown by his dreams, never reflects on his past pleasures or pains in the chase?” How indeed?
Culture: Science & History
A female Boxer provides the DNA for the first complete sequence of the dog genome—what will it mean to the health of man and dog?
“The dog is everywhere what society makes him,” wrote Charles Dudley Warner in the January 1896 issue of Harper’s New Monthly Magazine. Elaine Ostrander and Heidi Parker update that message in the November 2005 issue of the online journal, Public Library of Science—Genetics: “The domestication of the dog from its wolf ancestors is perhaps the most complex genetic experiment in history, and certainly the most extensive.” Undeniably, the results of that experiment are directly manifest in the appearance and behavior of the dog.
Since hitching its evolutionary fate to that of humans some 9,000 canine generations ago, the dog has proven the most adaptable, versatile and steadfast of companions, serving as a guard; draft animal; hunter; herder; warrior; entertainer; finder of explosives, contraband, disease and lost souls; healer, therapist; physical and spiritual guide; and friend. With the public unveiling last month of the fully sequenced and richly annotated dog genome—the approximately 2.4 billion base pairs of DNA (A [adenine], which always binds with T [thymine], and C [cytosine], which binds with G [guanine]) that form its genetic code—the dog might now also add to its monikers, shall we say, “genomic consort.”
Of course, the dog is not the first mammal other than humans to have its genome fully sequenced—the mouse, rat and chimp got theirs first—but because of its architectonic breed structure, it might prove the most illuminating. To shift metaphors: geneticists can now use the dog genome sequence like a combination zoom and telephoto lens, zeroing in on specific genes and even minute changes within genes, or jetting back to examine broad patterns and interrelationships within it and between it and other genomes that reveal the evolutionary history of an individual, a breed, a population, or the entire species and genus.
Dog as Cultural Construct
By most estimates, there are today more than 400 breeds worldwide, many with specialized morphologies and behaviors, nearly all genetically isolated. The majority of those breeds are also susceptible to one or more of more than 400 genetic disorders, approximately 350 of which are also found in humans, including epilepsy, kidney cancer, deafness, blindness, auto-immune disorders, congenital heart disease, skeletal malformations, neurological abnormalities, bleeding disorders and neuropsychiatric disorders.
Because traits and diseases often cluster according to breed and because breeders maintain extensive pedigrees, canine geneticists have long argued that the dog represents an ideal natural model for examining how genes shape appearance, function, behavior and health. In 1991, Jasper Rine, a geneticist at the University of California, Berkeley, with two researchers in his lab—Mark Neff, a postdoctoral fellow, and Elaine Ostrander, a staff scientist—started the Dog Genome Project to study those issues and in the process create a map of the dog genome they could link to that of the human genome for comparative study. [Ed. Note: See interview with Mark Neff.] From that effort was born a cottage industry, under the informal leadership of Dr. Ostrander (now at the National Human Genome Research Institute of the National Institutes of Health) and involving a small group of scientists worldwide, devoted to sequencing the dog genome, segment by segment. The researchers also lobbied to have the dog genome sequenced as part of the continuing Human Genome Project, in order to complete the task quickly and accurately.
In 2003, scientists at the Institute for Genomic Research and what is now the J. Craig Venter Institute in Rockville, Md., published a proprietary sequence covering 75 percent of the genome of a Standard Poodle, Shadow. On the other hand, sequences prepared as part of the Human Genome Project are posted in public data banks in the US, Europe and Japan as soon as possible after they are completed, so researchers can have access to them.
In July 2004, without fanfare, researchers from the Broad Institute at MIT and Harvard and Agencourt Bioscience Corp., of Beverly, Mass., led by Kerstin Lindblad-Toh, deposited in those public data banks their “first draft” sequence of 98 percent of the genetic code for “dog” in general and Tasha, an inbred Boxer from upstate New York, in particular. Their sequence was more complete and considerably more detailed than that of Shadow.
Then came a pause—of the seemingly interminable sort that occurs between the time certain dogs are called while snorfling in the park and the time they decide to respond—that was devoted to revision, analysis and assigning chunks of the sequence to their appropriate chromosome. The dog has 38 pairs of autosomal chromosomes—inheriting one from each parent—and two sex chromosomes. Lindblad-Toh’s team did not sequence the Y (male) chromosome.
Finally, at a press conference in Boston on December 7, 2005; in a lengthy article in the prestigious journal Nature on December 8; and in supplemental articles in the December issue of Genome Research, Lindblad-Toh and her team, along with Ostrander and dog genome scientists, officially unveiled the by then 99 percent complete sequence of Tasha and a SNP (pronounced “snip”) map showing 2.5 million “single nucleotide polymorphisms,” or mutations, in the genomes of Tasha, nine other purebred dogs, four wolves and a coyote. This map is useful for finding genes and examining interrelations between groups and individuals. The Nature article also contained an analysis by Lindblad-Toh’s research team of the dog genome’s structure and a new look at the dog’s family tree, origins and transformation by humans. (Full disclosure: Bark deadlines being what they are, I did not attend the press conference, which was designed to receive maximum coverage in the daily media.)
The Dog Genome
From a certain perspective, the dog is just another mammal, albeit with a genome slightly smaller and “cleaner”—“there is less junk,” Lindblad-Toh said—than that of its human companion or the ubiquitous lab rat, to which the researchers also compared it in Nature. Tucked within the 2.4 billion base pairs of the dog’s DNA are some 19,300 genes. By comparison, the human genome consists of approximately 2.9 billion base pairs of DNA and, at most recent count, approximately 22,000 genes. Approximately 72 percent of the dog genes are orthologous, meaning they correspond on a one-to-one basis with genes found in the human and rat genomes, although their functions might differ.
Comparison of mouse, human and dog genomes have identified a core 812,000,000 base pairs (5.3% of the total human genome) of ancestral sequence common to all three species. This DNA encodes proteins (1-2% of the total genome), and includes specific sequences that control gene expression. This portion of the genome is under what biologists call purifying selection, wherein variations on a gene or changes in a sequence are selected against, or weeded out. The sequencing of additional mammalian genomes, including those of Rhesus monkey, cow, opossum, elephant, rabbit, cat and shrew, should help to sharpen the focus on the DNA definition of mammal-ness.
Despite the similarities between all three species, it appears the genome better reflects the social reality of dogs and humans than does taxonomy, which places the rat closer evolutionarily to humans than the dog. The researchers reported in Nature that some sets of functional genes, like those involved in brain development, showed signs of having evolved similarly in dogs and humans—and more rapidly than in rats. It is a suggestive finding.
The dog’s value in comparative genomics lies in large measure in its breed structure, and here the researchers offer some support to a couple of recent suggestions that repetitive segments of DNA are somehow tied to the dog’s physical plasticity, its ability to assume so many different shapes and sizes, as well as to fall victim to various inherited diseases. Geneticists have focused not only on SNPs—changes in a single base—but also on repetitive blocks of DNA, including “short interspersed nuclear elements” and “tandem repeats.”
SINE elements, as they are known, are repetitive segments of DNA between 150 to 750 bases long. Interspersed throughout a genome, they move around over time, and some are species-specific.
On the whole, the dog genome has fewer SINE elements than the rat or human. But it has a “highly active carnivore-specific SINE family” that is full of mutations that vary between breeds, Lindblad-Toh and her co-authors wrote in Nature. These SINE elements are greater in frequency by a factor of at least 10 than any found in humans, and are believed to play a role in gene expression. When inserted into genes, they can cause diseases, like narcolepsy in Doberman Pinschers and centronuclear myopathy (a muscle disease) in Labrador Retrievers.
Wei Wang and Ewen F. Kirkness of the Institute for Genomc Research, writing in Genome Research, argue that SINE elements are a major source of genetic diversity in the dog. Citing their research, Lindblad-Toh and her colleagues speculate in Nature that the variation from SINE elements “has provided important raw material for the selective breeding programs that have produced the wide phenotypic variations among modern breeds.” In that event, SINE elements may have been what has allowed humans to produce everything from the Pug to the Irish Wolfhound.
But no one knows. In December 2004, John W. Fondon, III, and Harold R. Garner of the University of Texas Southwestern Medical Center proposed in the Proceedings of the National Academy of Sciences—in a paper that caught the attention of scientists, if not the press—that changes in the length of “tandem repeats” found within genes are responsible for the phenotypic variation between breeds and the speed with which breeders can change a breed’s appearance. Once called “junk DNA,” like so many other parts of the sequence whose purpose was then unknown, these “tandem repeats” occur when two or more nucleotides form a pattern that repeats itself over a short stretch of the genome.
Simply, tandem repeats are shorter than SINE elements. Both are suspected of playing a role in creating the plethora of dog breeds. But changes in the timing of development are also believed to be involved. Sorting that out is where the Dog Genome Project began and where it still must go. In that sense, the genome sequence represents a beginning rather than an end.
A Genomic Look at History
It is thus poetically fitting and perhaps scientifically significant that genetically, dogs show a similar pattern of homogeneity and variability between and within breeds, especially the modern breeds. In general, they were formed through extensive inbreeding and the use of favored sires, both of which serve to limit genetic diversity. Yet, despite that, or perhaps because the genetic isolation of breeds has not been long enough or as extensive as breeders sometimes claim, those breeds continue to possess a surprising amount of genetic diversity.
It is easier to assemble the genomic sequence of highly inbreed animals because of the genetic homogeneity of the pairs of chromosomes being sequenced. Compared to other Boxers, Tasha’s genome has one SNP—one change in one letter, or nucleotide – every 1,600 base pairs. Less inbred breeds would have more SNPs. Such changes or mutations appear randomly throughout the genomes of all animals, primarily in non-coding regions outside genes, where their purpose, if any, is uncertain, and far less frequently within genes where they can cause lethal mutations.
SNPs persist for hundreds of generations and form distinctive, inheritable clusters or blocks of genetic code on chromosomes that are known as “haplotypes.” Because they are passed on through generations, haplotypes are useful for exploring the evolutionary history of individuals, groups and species, and seeking out clusters of genes involved in inherited diseases, in morphology and, it is hoped, behavior. Probing the differences between individuals with congenital heart disease, for example, and those without, researchers would use their SNP map to identify the haplotypes of sufferers against those who are disease-free in an effort to find a region or regions on a chromosome that seemed involved. There, they would focus the search for genes.
To create a densely detailed SNPs map of the dog genome, Dr. Lindblad-Toh’s team partially sequenced the genomes of nine additional dog breeds, four kinds of wolf, and a coyote: German Shepherd, Rottweiler, Bedlington Terrier, Beagle, Labrador Retriever, English Shepherd, Italian Greyhound, Alaskan Malamute, Portuguese Water Dog, Chinese gray wolf, Alaskan gray wolf, Indian gray wolf, Spanish gray wolf, and California coyote. They also had the genome sequence from the French Poodle, Shadow. The researchers found 2.5 million cases where there were differences in a single nucleotide between the various canine genomes.
Comparing the Boxer to the other breeds and the five wild canids, the researchers found that while the SNP rate between different Boxers was 1 for every 1,600 base pairs, it was around 1 for every 900 base pairs between the Boxer and every other breed but the Malamute, which was 1 for every 787 base pairs. According to the breed identification system developed by Elaine Ostrander and Leonid Kruglyak of the Fred Hutchinson Cancer Research Center in Seattle, the Malamute belongs to a class of “ancient dogs” and thus would be expected to be more distant from the modern Boxer than other modern breeds—the shorter the distance between SNPs, the more distant the relationship. The other dogs represented breeds created or consolidated within the past 300 years, like the Boxer. Compared with the Boxer, the wolves had ratios of around 1 SNP for every 580 base pairs, and the more distantly related coyote stood at 1 for every 420 base pairs.
The Boxer’s genome represents “a mosaic of long, alternating regions of near-total homozygosity and high heterozygosity,” the researchers reported in Nature. The homozygous regions, wherein both chromosomes in a pair have identical haplotypes, cover 62 percent of the genome; the heterozygous regions, in which the haplotypes are not identical due to SNPs or genetic variations, 38 percent.
The researchers then scanned the genomes of 20 dogs from each of 10 other breeds, and one dog from each of 24 breeds, ranging from the ubiquitous Labrador and Golden Retrievers to the rare Glen of Imaal Terrier, but always limiting themselves to purebred dogs registered by the American Kennel Club. The analysis by Lindblad-Toh’s team reported in Nature that most dog breeds were similar to the Boxer in terms of number of SNPs, and the relative proportions of heterozygosity and homozygosity were also similar.
The one aberration was the Akita, a Japanese breed created, by Lindblad-Toh’s estimate, some 10,000 years ago for hunting, which passed through a bottleneck in the 1940s in America. The first official Akita in America was a gift from Japan to Helen Keller in 1937.
Using mathematical models that postulated an effective population of 13,000 dogs with an inbreeding coefficient of .12—meaning basically that they are cousins—the researchers concluded that in achieving its current blend of sameness and difference, the dog passed through a major genetic bottleneck 9,000 generations ago, and another 30 to 90 (sometimes given as 50 to 100) generations ago. Assuming a generation time of three years for dogs, they pegged the origin of the dog at 27,000 years ago from perhaps as few as two wolves. Lindblad-Toh said in an e-mail that the founding population of wolves might have been larger—and some geneticists say there must have been several hundred animals involved—but the genome does not appear to record any contribution from them. She also said that there may have been multiple domestication events and back-crosses with wolves at various times and places, as other genetic studies have shown.
I am a fan of ancient dates when it comes to dog origins, and the older the better, but this new offering—a sort of compromise between 15,000 years ago in East Asia, proposed by Peter Savolainen of the Royal Institute of Biotechnology in Stockholm and his colleagues in 2002, and 40,000 to 135,000 years ago that Robert K. Wayne and his lab team at the University of California, Los Angeles (including Savolainen), proposed in 1997.
Geneticists are divided, but archaeologists are not. Darcy Franklin Morey, an archaeologist at the University of Kansas specializing in the dog, says that 27,000 years ago, like the older dates, fails to coincide with the archaeological record, which dates to around 12,000 to 14,000 years. Morey has a paper forthcoming in the Journal of Archaeological Science arguing that the proliferation of dog burials at the time marks the origin of the dog. He does have a point in that as a cultural construct, the dog has left natural history and entered human history. Arguably, then, the molecular clocks used to calibrate its age should be set to human, as well as geologic time. Beyond that, the choice of three years as the generation time for dogs is unexplained and possibly long.
A notable change in the dog from the wolf is that females enter first estrus at six months to one year of age. Breeders in the rural South have long bred their dogs at first heat, and it may well be a custom dating back centuries. Australian dingoes, living in packs independent of men, start breeding around two years of age, according to dingo expert Laurie Corbett in The Dingo in Australia and Asia. Beyond that, my searches fail to pull down any calculation for the generation time for dogs. Clearly, that needs more examination.
In a commentary accompanying Nature’s presentation of the genome, Hans Ellegren, an evolutionary biologist at Uppsala University, raised another qualifier, saying that if “repeated back-crossing has occurred” between dogs and wolves, Lindblad-Toh’s model of the dog’s origins “would have to be revised.”
The Broad Institute team’s analysis appears on firmer footing when it comes to placing breed formation within the past 200 to 300 years. Breeds are formed through consolidation of working dogs of an existing type into a more consistent form, through hybridization and inbreeding to reconstruct a breed, and through building on a small number of imported animals. Early dogs bred rather freely, and—because in sexual reproduction, the chromosomes from the sire and the dam are recombined to form the single chromosome passed on from each and because alignments are not perfect—haplotypes were broken up, becoming shorter and more scattered over the generations of random breeding.
Through inbreeding from a small gene pool to create their particular breed, humans unknowingly selected a small group of overlapping chromosomes carrying the genes for the traits they wanted—and for some diseases they didn’t want. From that came the breed’s distinctive pattern of large homogeneous haplotype blocks and shorter heterogeneous ones, which selective breeding has sustained.
A notable exception to this model appears to be the Labrador Retriever, which replaced the “old yellow dog,” or cur dog, as America’s most abundant big dog and so is less inbred than other breeds, except in some lines. The English Springer Spaniel and Golden Retriever are more inbred than Labs but less than most other breeds. But all three show the pronounced influence of “favorite sires,” whose overuse also serves to limit diversity. In an interview, Elaine Ostrander said research in her lab has indicated a clear genetic break between show Labs and field-trial Labs, and between show and hunting Beagles.
Breeds that passed through a formation bottleneck typically have four of around 10 possible haplotypes, the researchers said. Although the haplotypes and their proportions vary from breed to breed, haplotypes are also shared between them. As the Akita indicates, this arrangement might not extend beyond breeds of western dogs. But it means that researchers can use 10,000 SNPs, a relatively small number, to scan the genomes of dogs within the breed and compare them with those of other dogs in the same breed in order to find genes.
“We now have a whole new tool kit for looking at the evolutionary history of canines and the origins of dogs, where they originated and how they spread, and how often they interbred with local wolf populations,” said Robert K. Wayne, an evolutionary biologist at the University of California, Los Angeles. A 60,000-SNP GeneChip (similar to a computer chip but incorporating DNA) soon to be available from Affymetrix should speed the search for genes that regulate a dog’s phenotype, he added.
Wayne and his lab contributed a new family tree, or phylogeny, of 34 canid species for the Nature paper, showing the time of their emergence during the past 40 million years and solidifying the already strong argument that the wolf is the dog’s nearest relative, while the wolf’s wild kin are, in order, the coyote, golden jackal, Ethiopian wolf, dhole and African wild dog.
A small group of scientists believes that one of the dog’s more distant relatives, the fox, specifically a colony of tame foxes in Siberia, might hold the key to the genetic changes underlying domestication. They are using the dog genome, said one of the leaders in that quest, Gregory M. Acland, a geneticist with the James A. Baker Institute for Animal Health at Cornell. Like a number of other geneticists, Acland predicts that within 20 years, SNP maps will be rendered obsolete as chips and programs are developed that allow the entire genome of an individual, or parts thereof, to be sequenced cheaply and quickly.
He likens genomics to a sophisticated video game. “You start playing this game,” he said, “killing everything you see or collecting things, and after you’ve killed and collected everything there is and the game seems over, a little box appears in an upper corner. You click on it, and suddenly you’re in a whole new, more complicated level.”
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