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Good Dog: Studies & Research
The "D" Word: Dominance
We talk about dominance, but do we really understand it?

If a dog has behavior issues such as a tendency to mount other dogs, any form of aggression, an overly pushy play style or poor response to training, some people are sure to claim that “dominance” is the culprit. But are they right?

The ongoing dialogue about dominance in the dog world is more problematic than an unattended puppy in a shoe store but it’s unlikely to go away anytime soon. While some hate the concept so much that they refer to it as the “D-word,” others swear by it, considering it an indispensable guiding principle for all interactions with dogs. Having a civil discussion on the subject, much less reaching a consensus, is a challenge, as debates often become quite heated.

Scientific inquiry offers an opportunity for greater understanding of this topic, though writings on the subject with titles such as “Social dominance: Useful construct or quagmire?,” “Social dominance is not a myth: Wolves, dogs, and other animals” and “Deconstructing the concept of dominance: Should we revive the concept of dominance in dogs?” reflect continued controversy. While dominance, or social dominance as it is often called, has been studied extensively in a number of species, relatively little work has been done in this area on the domestic dog, and more research is badly needed.

Defining Dominance
To further complicate an already contentious debate, “dominance” has multiple meanings. As James O’Heare notes, “Often, researchers argue over social dominance without agreeing on a clear definition, and perhaps have different definitions in mind.” There are a variety of ways to characterize it. In his study “The concept and definition of dominance in animal behaviour,” Carlos Drews reviewed 13 different definitions of dominance and pointed out that criticisms of one are not necessarily applicable to others.

Adding to the frustration and confusion, when it comes to domestic dogs the term is commonly applied to two different types of relationships. The first relates to interactions between dogs. In this usage, dominance is defined as the power to control access to desirable resources and refers to the relative status of two dogs. In the absence of another dog, an individual dog cannot be said to be “dominant” as a personality attribute because dominance refers to the relationship between dogs.

The second, and more controversial, type of dominance relationship relates to interactions between humans and dogs. In this paradigm, humans dominating dogs is considered the path to well-trained dogs. Those who follow this school of thought claim that you have to control your dogs by being dominant over them in order to make them behave and may make suggestions such as not allowing your dog to sleep on your bed or walk through the door ahead of you, or even to spitting in your dog’s food and making a resting dog move rather than walking around him. Today, fewer trainers subscribe to these ideas than in the past.

Dominance Between Dogs
The few scholarly investigations of dominance relationships between dogs have found the concept relevant to their interactions with one another, and scientists who study social relationships aren’t surprised. For example, Marc Bekoff, PhD — ethologist and professor emeritus at the University of Colorado who has had a major influence on the scientific understanding of dogs as well as many other species — observed that “in the real world, dogs are social beings, and social beings form dominance relationships, just as their wild relatives do.”

Becky Trisko, PhD, ethologist and owner of Unleashed in Evanston, Ill., focused her 2011 doctoral dissertation on social interactions within a group of 24 dogs who regularly engaged with one another at daycare. The behaviors she analyzed included aggressive threat and conflict, muzzle lick, crouch, passive submission, retreat, high posture, muzzle bite, mount and chin-over.

Trisko found a dominance hierarchy among the dogs, although only about 30 percent of the pairs had clear dominance relationships. Dominance rank correlated with age (older dogs tended to rank more highly) but not with size. And contrary to popular belief, neither mounting nor performing chinovers were related to status. As a point of interest, not once in 224 hours of observation during this yearlong study did she observe an “alpha roll.”

Muzzle licking was consistently done by subordinate dogs to individuals of higher rank and was highly predictive of relationships between individuals. The clearest signals were those associated with voluntary submission, or deference. Trisko observed that dominance relationships were not about coercion, force or fighting, but rather, about an understanding by both individuals of their relative social status.

In another study exploring dominance relationships between dogs, Simona Cafazzo and three colleagues observed a group of feral dogs in the suburbs of Rome. Their primary finding was that the dogs formed a linear dominance hierarchy, meaning that the individuals in the group could be ranked in order from highest to lowest in status. (Other possible social structures include having one individual who dominates all others who are equally low-ranking or societies in which the relationships are not transitive— e.g. A dominates B, B dominates C, C dominates A). A linear dominance hierarchy indicates that the dogs in this study were capable of forming stable social groups, although many have claimed that feral dogs cannot do so. Additionally, they found that submissive behavior was most predictive of dominance relationships, rank correlated with age, and males within an age class outranked females. Rank order in the linear dominance hierarchy predicted access to food resources, with those of higher rank having priority access.

Dominance has been studied in puppies as well as in adult dogs. John Bradshaw and Helen Nott reported that interactions between littermates were inconsistent over time, and that observations of such interactions did not predict which puppy would come out on top in any competitive situation; “winners” varied from one day to the next. Despite much discussion of choosing (or avoiding) the dominant puppy in a litter, interactions between littermates do not reveal dominance relationships, much less any kind of linear hierarchy.

The Dangers of Misunderstanding Dominance
Conversations about dominance relationships between dogs are often tense, but discussions of the concept within the context of relationships between humans and dogs are sometimes nothing short of explosive. Many of the strongest objections stem from using it to endorse force-based, coercive styles of training. According to Trisko, “Dominance has been wrongly equated with aggression and used to rationalize the use of physical force and intimidation by humans toward dogs. Misunderstandings of the concept of dominance have led to unnecessary physical punishments and abuse of dogs by humans.”

While no studies have thoroughly investigated whether dominance relationships exist between people and dogs, there is evidence that such training styles can create problems. Herron et al. investigated such techniques, often called “dominance reduction training,” and found them to be counterproductive. The confrontational methods associated with training styles that insist that we “get dominance over our dogs” caused aggressive responses in 25 percent of the dogs in their study. Techniques such as grabbing a dog by the jowls and shaking; hitting or kicking; staring; performing alpha rolls (also called “dominance downs”) and physically forcing a dog to release an item were more likely to result in aggressive behavior than were gentler, positive methods. Using such forceful methods can actually create problem behavior as well as increase a dog’s fear and anxiety. Based on what we know about dominance relationships between dogs, this is not surprising. As Trisko notes, “If dominance relationships between dogs and humans are at all similar to dominance relationships between dogs, then dominance does not apply to all relationships and when it does apply, it does not require the use of intimidation or physical force.”

A basic ethological premise is that we must understand the animals we study. In fact, this principle is considered so absolute that it is most often phrased as a commandment: Know thy animal! Understanding how social dominance does and does not apply to dogs is part of knowing who dogs are. Trisko makes this point: “If we really want to understand our dogs’ behavior, especially their relationships with other dogs, ignoring dominance will hinder us.” Bekoff agrees. “That’s who they are, that’s how they behave. They form status relationships, and we have to understand that.”

When it comes to the issue of dominance, common ground is not easy to find. Few would dispute the need for further research, though even the most carefully designed studies may not be enough to bring agreement on this particular subject. As Bekoff has noted, “People get on this kick with dominance. They don’t pay attention to the data.”

Arguments about dominance and its relevance to dogs, their relationships with each other, and our relationships with them are sure to continue. Though I prefer resolution to conflict, I can’t help but see the wisdom in moralist and essayist Joseph Joubert’s remark: “It is better to debate a question without settling it than to settle a question without debating it."

Culture: DogPatch
Are Dogs’ Mouths Cleaner Than Humans’?
Young Scientist in Action
Abby Walling & Lucy

Are dogs’ mouths cleaner than humans’? Her grandmother said they were, her mother said they definitely were not. Not one to take things at face value, Abby Walling of Iowa City, Iowa, decided to conduct an experiment and answer the question once and for all. Not only did she reach a conclusion, she won a 2011 Young Naturalist Award from New York’s American Museum of Natural History for her work. As a bonus, she no longer feels guilty about letting Lucy, her kiss-happy Yorkshire Terrier, lick her face.

Specifically, Abby investigated the bacterial content of both human and canine tongues. She hypothesized that humans would have fewer bacteria — after all, they brush their teeth regularly and aren’t quite as indiscriminate as dogs about what they put in their mouths. She then obtained saliva samples from five dogs and their associated humans and plated the samples on agar in a laboratory. What she found surprised her: Humans do have more bacteria in their mouths than dogs. That, however, is only part of the story (to read Abby’s full project report and results, go to amnh.org/yna and look under Past Winners, 2011).

At the time, Abby was in the eighth grade at Northwest Junior High School, and this project landed her a place in the State Hygienic Lab’s student mentorship program, which matches junior high and high school students with a laboratory scientist mentor; Gabe Gerken, public health microbiologist, helped her refine her methodology and the lab provided testing supplies.

What does Abby see in her future? “After college, I’d like either to manage a zoo or conduct scientific research concerning animals and the environment. I am very passionate about improving the world, and I believe I can use research to solve some of the environmental problems the world is facing today.”

Sounds like we’re in good hands.

Good Dog: Studies & Research
Education: Gone to the Dogs
Canines claim their territory on college campuses

Summer has faded into fall and it’s time for dog lovers — and dogs too — to head to college, where dogs are taking their place in the dorm, the psych lab and even the classroom.

While some dogs simply kick back and enjoy campus life at a university with pet-friendly housing, such as Eckerd College in St. Petersburg, Fla., or Stephens College in Columbia, Mo., others give their intellectual muscles a workout by participating in research studies designed to test their ability to think and solve problems.

Dogs are taking their place in collegelevel human studies as well. At several universities, they dominate the syllabi of courses devoted to companion-animal behavior and welfare. Other schools offer entire classes or majors focused on the human-canine connection.

To a dog lover, the appeal of taking a dog to college is obvious, as is the draw of canine-focused study — but what’s in it for the dogs? While it has long been possible to study animal science, wildlife management or food-animal husbandry, formal study of dogs in academia is a relatively new phenomenon. As recently as the 1990s, academic researchers who wanted to focus on Canis lupus familiaris were greeted with raised eyebrows, ridicule or worse. But the nascent fields of anthrozoology — the study of human-animal relationships — and cynology — the study of the domestic dog — are growing quickly in academia.

Applying academic rigor to the study of dogs can increase our understanding of their abilities and deepen our bond with them, ultimately resulting in improving their treatment by society as a whole. These goals spurred the development of the country’s first anthrozoology program and the establishment of a university wholly devoted to the study of dogs. Both of these pioneering efforts are part of a growing collection of canine-focused educational options.

When Anne Perkins was head of the psychology department at Carroll College in Helena, Mont., she was dissatisfied with existing animal-focused study options, which were basically limited to animal science and zoology. “These programs were not addressing why we love our animals so much,” she says. Perkins spent a 2005 sabbatical designing a new program, anthrozoology, which would “study the value of animals from an academic, scholarly perspective.” The new program was first offered as a minor at Carroll in 2007.

“I bit it off in pieces,” she says, adding one class at a time. The students wanted more; the minor grew into a major, and Carroll offered the nation’s first bachelor of arts in anthrozoology in 2011. The bachelor’s degree “embedded the study [of the animal-human bond] in traditional fields,” where research is peer-reviewed and published in scholarly journals, Perkins says.

Carroll College anthrozoology students focus on either horses or dogs. Students in the canine track examine theories of domestication and attachment. They study puppy development, socialization and learning, and they practice assessing temperament. Seniors raise puppies, preparing them for a broad range of doggie careers, including scent work, assistance and acting.

Like Perkins, Bonita Bergin, founder and president of Bergin University of Canine Studies (BUCS), argues that academic study is essential to improving the status and treatment of dogs. As BUCS graduates leave the Rohnert Park, Calif., campus to teach or run businesses that model ethical humancanine relationships, “we hope to enrich the understanding of the relationship that has inspired and fulfilled so many,” Bergin says. “We also hope to help eradicate the horror of euthanasia of unwanted dogs.” Offering post-secondary study wasn’t enough for Bergin; she also wanted the respect of academic peers. Tenacious as a terrier, Bergin spent three years pursuing her vision: the world’s first accredited university focusing on our canine pals and partners. Why? “I believed the dog deserved it,” she says simply.

Undergraduate and graduate students at BUCS explore the influence of genetics and heredity on dogs’ behavior and temperament. They also analyze the growing body of published research on dogs, and are encouraged to contribute original research of their own. But it’s not all books and theory. Puppies and service-dogs-in-training fill the campus with hands-on opportunities. The associate degree program, in particular, emphasizes dog training and socialization; starting the day students help out with the whelping process.

Bergin has revolutionized earlypuppy education. BUCS students begin “formally” training puppies as soon as the puppies open their eyes at about four weeks of age. The astonishing result is that most puppies respond eagerly and accurately to more than a dozen verbal cues by the time they are eight weeks old.

On the opposite side of the country, dog-loving students at SUNY Cobleskill choose among a half-dozen dog-focused electives in the animal science department. “[The courses] are designed to give students a solid understanding of the important factors involved in producing good working dogs and the behavioral basis of popular training techniques, emphasizing positive, reward-based approaches,” says Stephen Mackenzie, professor of animal science at the university. According to Mackenzie, a canine management major is in the works. Dogloving students “can work dogs almost every semester they are here,” he adds, training dogs for anything from offleash obedience and agility to tracking, trailing, air scenting and detector work “under the guidance of someone with good academic credentials.”

At some universities, dog scholars have to search for dog-related material buried like treasured bones among more traditional offerings. The psychology department at the University of Michigan, for example, offers “Dog Cognition, Behavior and Welfare,” a popular course taught by Camille Ward. The class, described as “for people who love dogs and want to learn about them from many different avenues,” has a long waiting list. Also in the psychology department, Dr. Barbara Smuts teaches “Behavior of Wolves & Dogs”; she also offers students the opportunity to participate in research projects on dogs’ social behavior.

At Barnard College, in New York City, Dr. Alexandra Horowitz (author of Inside of a Dog) teaches a psychology class on canine cognition. At Eckerd College, a course on animal learning and training includes considerable material on dogs, says its instructor, Lauren Highfill. The Center for the Human-Animal Bond at Purdue University in West Lafayette, Ind., offers companion-animal welfare and management courses that primarily focus on dogs and cats. Graduate students can head to Tufts University for a master’s program in animals in public policy that includes study of companion animals, or to Harvard, where psychology grad students can take a seminar called “Puzzles of the Mind: Humans, Animals, Robots.”

At Canisius College in Buffalo, N.Y., dogs figure prominently in undergraduate coursework on the social organization of animals, animal learning and applied animal behavior. And the college’s master of anthrozoology coursework includes a popular class on companion animals in society. A dog-human relationships expert was recently hired, and Canisius plans to expand its dog-centered offerings, says Michael Noonan, professor of animal behavior, ecology and conservation.

We’ve come far since 17th-century philosopher René Descartes asserted that animals lacked the ability to feel pain, yet cruel treatment of dogs is still far too common. Canisius prepares animal-behavior graduates to eradicate that cruelty and to “make the world a better place in the way we interact with animals” by providing a “strong, science- based education balanced with critical thinking and ethics,” Noonan says. “From the science, we see that they [animals] are more like us than was thought in the past.” Therefore, “most ethics that apply to us apply to them — animals are sentient beings whose concerns matter.”

Some schools recognize the importance of the human-animal bond by allowing pets in selected on-campus housing units — about a dozen colleges nationwide have at least one pet-friendly dorm. Other schools conduct research studies that aim to improve understanding of dogs’ abilities and view of the world. Indeed, new evidence of dogs’ intelligence, creativity and ability to understand and communicate their concerns is uncovered daily at cognition labs, where dogs take center stage.

New York City dogs can join cognition studies in Horowitz’s lab at Barnard where anthropomorphic beliefs about dogs are tested with an emphasis on “getting the dog’s perspective,” rather than a more traditional behavior-focused approach, said researcher and Bark contributing editor Julie Hecht. Current studies examine dogs’ understanding of the concept of “fairness” and the way they use their noses in daily life. “We’re trying to better understand the dog’s perspective, but we are, of course, limited by our human perspective,” and sometimes the hardest part is separating the two, she said.

Southern dogs have a choice of schools: Duke University (Durham, N.C.), the University of Florida (Gainesville), the University of Kentucky and Eckerd College all recruit local canine “students” for their research. Current studies examine whether dogs can count, how dogs form trusting relationships with humans, dogs’ interpretation of human social gestures, and canine imitation and social learning.

The studies might sound esoteric, but they can lead to real changes in the way people regard and teach dogs: Watching four-week-old puppies learn to sit, lie down and solve problems banishes forever any idea that training must involve force. Discovering that dogs can use pictures to indicate their preferences compels scientists to reexamine human-centered ideas that tie thinking to spoken language. And seeing how dogs’ behavior changes when they know that human “observers” are distracted hints at their ability to strategize.

The more we learn about dogs’ abilities, the greater the potential for true partnerships based on mutual respect rather than compulsion, says Bergin. “This is crucial in transitioning the dog from a backyard animal we see as disposable to recognizing the key role dogs play in the evolution and continued development of humans.”

Good Dog: Studies & Research
Dog Training: Operant Conditioning
When operant conditioning clicked (and clucked)

On a warm and slightly overcast morning in 1967, a rusty, mustard-colored station wagon slowly approached the terminal at San Francisco International Airport. Wheels still rolling, a door opened and something gray jumped out. As the wagon continued on its way, an animal headed toward the terminal. It was a cat.

Straight five steps, then wait. The glass door opened and as a portly man in a business suit dragged his overnight bag through it, the cat darted in. Straight 10 paces and the cat was inside the terminal. It headed left 20 feet, then right 30 feet, then left two more feet. No one seemed to notice. The cat settled under a bench where two men sat, engaged in intense conversation. Ten minutes passed, then 20; the cat lay patiently, its tail occasionally twitching.

Then, abruptly, the cat stood up and retraced its steps. Two feet to the right, 30 feet to the left, 20 feet to the right and out the glass doors. Once again, the station wagon pulled up and without stopping, a door opened. The cat leaped in. Mission accomplished.

The project, commissioned by the CIA and run by Animal Behavior Enterprises, had been a success. The cat’s cochlear implant (a device agents used to listen to the cat’s environment) had proven reliable, and its months of training using the relatively new technology of operant conditioning had proven effective for this intelligence operation.

Does this sound preposterous? Would it sound less preposterous if the trained animal had been a dog? Thanks to the science of operant conditioning, European police and military teams have been able to train their working dogs to perform at a much higher and more reliable level than had been possible using traditional coercion- based methods.

This type of training is no small feat. In 1996, Simon Prins, co-author of K9 Behavior Basics: A Manual for Proven Success in Operational Service Dog Training (2010), was hired to lead an innovative project for the Canine Department of the Netherlands National Police Agency. A test project with a three-year timeline, it would continue if it were a success. The project included a detailed list of tasks for dogs to perform.

“This included normal operational tasks, such as tracking, and explosive and narcotics detection,” says Prins, “but also climbing, rappelling, traveling by helicopter and boat and, the most challenging, training dogs to work with cameras and to follow radio or laser guidance at long distances.”

Although Prins had been a patroldog handler in the regional police force for only a few years, he was selected for this project because he was seen as an innovator. “I had been questioning our traditional force techniques because I noticed that dogs would shut down and stop working, or my police dog would become aggressive to me and to the trainer. So I was already looking for new methods.”

Stepping Up
New methods were what the job was all about. “The other traditional [forcebased] trainers all said that radio or laser guidance was not possible,” according to Prins. But he was sure there must be a way; so, even though he would not be able to return to his old job if he failed, he accepted the challenge. Within three years, he had succeeded in completing all of the tasks set for him, as well as a few more.

At this point, you may be asking yourself — given the fact that people have been training dogs for more than 4,000 years — why did traditional trainers feel these new tasks were impossible? Also, if a guidance system had already been developed for cats in 1967 in the U.S., why did it take Prins three years to reinvent the wheel 30 years later?

Bob Bailey, who worked on the 1967 project and later became co-owner of Animal Behavior Enterprises after marrying its cofounder, Marian Breland, explains. According to Bailey, it was the advent of animal training and behavior as a science that allowed them to develop the system for dogs, cats and, later, dolphins. “Dog training has been practiced as an ancient craft,” says Bailey. “The science of training wasn’t developed until the 1940s with B.F. Skinner.”

What’s the difference between craft and science? According to Bailey, “Crafts generally develop over thousands of years and tend to preserve what’s old and what has been done before. Information is passed down in secret from master to apprentice, and the apprentice must never question the master.” As a result, when errors are introduced, they tend to be preserved. Another characteristic of a craft is that a change is usually designed only to solve an immediate problem. “Rarely do they look for general principles,” says Bailey.

Science, on the other hand, is a systematic way of asking questions, a process that eventually weeds out mistakes. It’s guided by principles and data, and researcher’s approaches change and are revised as new information comes to light. As a result, science advances quickly compared to craft.

Bailey backs up his description with an example: “For 1,000 years, the Chinese used gunpowder to build small rockets. Then the Turks decided to build bigger ones, which they used on the British. It took them 800 years to develop the technology.” Then, in the 1900s, science and technology stepped in. In 1926, American rocket pioneer Robert Goddard launched the first liquid- propellant rocket. In 1949, less than 25 years later, the U.S. sent a rocket to the moon.

“So it took 800 years of craft to send a six-foot rocket half a mile and less than 50 years of science to send a rocket to the moon,” Bailey summarizes.

From Puzzle Boxes to Bells and Whistles
Once the psychology of learning became a science, animal training made the same great strides. In 1898, scientist Edward Thorndike first described the principle of operant conditioning. Using cats and a small cage called a puzzle box, with food outside the box, he found that cats placed in the cage could learn how to get out, and that with each trial, they got faster at it. He theorized that the cats were learning by trial and error.

Others were making discoveries at the same time, but the one who really put things together was B.F. Skinner. Through many experiments, Skinner developed the principles of operant conditioning, which describes how animals cope with their environments.

Skinner found that animals learn to repeat behaviors with consequences they view as positive and to avoid behaviors with those they view as negative. They learn best when the positive or negative consequence is timed exactly to the behavior, and their learning rate is directly proportional to the rate at which the behavior is reinforced.

Research and technology advanced quickly, and within only eight years, operant conditioning had made its way out of the lab into an applied setting. During World War II, Skinner, who wanted to help with the war effort, set out to train pigeons to guide missiles by pecking at an image of the target site on a screen in a project called Project Pigeon. To demonstrate to the navy how it worked, Skinner took six pigeons and the apparatus in which they were trained to Washington, D.C. The demonstration was successful, but the navy turned him down; the admirals may have been taken aback when Skinner opened the chamber, which resembled a Pelican missile warhead, and they saw three pigeons pecking away.

Skinner’s two graduate students, Marian Breland and her husband, Keller Breland, had dropped out of their degree programs to help with Project Pigeon. They learned a lot while working with Skinner, more than they had learned in class, and became skilled at shaping — a process by which a goal behavior is taught in increments, systematically rewarding intermediate behaviors. They also learned about secondary reinforcers — unique tones such as a clicker or whistle that, when paired with food, could be used to tell the animal exactly when it had done something right.

The couple, who saw how powerful these nonforce techniques could be, decided that when the war was over, they would get into some kind of business where they would use operant technology to help solve problems for animals and, later, humans. In 1947, they founded Animal Behavior Enterprises (ABE), whose mission was to demonstrate a better, more scientific way of training animals in a humane manner using less aversives.

They started with dogs, but trainers shunned the new method, claiming that people had been successfully training for centuries and no new approaches were needed. Rebuffed on the canine front, the Brelands turned to other species. For 47 years, ABE trained animals for its own theme park, the IQ Zoo, in Hot Springs, Ark., as well as for shows across the country. At ABE’s height, the Brelands could have up to 1,000 animals in training at any given time, many for companies such as General Mills, who used them in commercials and at sales conferences. They also worked on animal behavior and training projects for the U.S. Navy and Purina, as well as for Marineland of Florida and Parrot Jungle, where they developed the first of the now-traditional dolphin and parrot shows. When they started, there was only one trained dolphin, whom it had taken trainers two years to get ready to perform. In six weeks, Keller trained two new dolphins to perform the same behaviors.

Bob Bailey met the Brelands when he was hired as director of dolphin training for the navy and Keller and Marian were contracted to help. “I spent six months at ABE learning to train many animals, including chickens.”

Three years later, the same year Keller passed away, Bailey joined ABE as assistant technical director and head of government programs. Later, he became research director too and then executive vice president and general manager. Eventually, he and Marian married.

Over the course of their career, the Baileys trained more than 140 species (or about 16,000 individual animals). In 1990, they retired and closed ABE. Then, in 1996, they received a series of calls from Simon Prins.

Inspired by Dolphins
“I’d spent a year traveling internationally and searching for training methods. I saw a dolphin show, and the girls were training the dolphins to perform incredible behaviors without shock collars or punishment devices. I did more research, and it all led back to Marian and Bob Bailey, so I contacted them,” Prins recalls. It took 20 emails and 10 phone calls before the couple agreed to help. “Bob was quite rude … he hung up on me many times.”

“We didn’t want to deal with police or military because in our experience, they are punishment-based,” says Bob. In the U.S., the Baileys had come to feel that force-based trainers could not make the change to operant technology because eventually, they fell back on the method with which they were most comfortable. “These trainers take what we say and modify it. They take good operant-conditioning principles and modify them, and then say they won’t work.”

Eventually, as Prins continued to meet Bob’s increasing demands, Bob agreed to help. Prins came with a few other trainers as well as his superiors to the Baileys’ Hot Springs headquarters to learn by training chickens.

Animal Behavior Enterprises had tested many animals for learning purposes and found that chickens provide by far the best training model (find out why). Prins and his bosses quickly learned that training is a technical skill rather than a mystical, inborn ability. A science, not a craft. They trained chickens to selectively peck just one type of object among a group of objects, and to perform tasks only on cue. They learned to train behaviors as a series of many little shaping steps, and to keep track of the outcome of each trial in order to determine whether they were having success or needed to fix their technique or plan. They did this all with positive reinforcement — without physically manipulating the chickens.

“Bob and Marian changed my whole perspective on animal training,” says Prins. As a result, he met all of the 1996 goals, and more. At first it was difficult. ABE had developed remote-guidance systems for cats, dogs and dolphins by 1967. In months, they could train dolphins to perform many behaviors, including traveling 12 hours on a circuitous eight-mile route with no reinforcement. It took Prins three years to work out the methods.

“I talked to Bob by mail and phone, but it was difficult, because I was the only one here using these techniques,” he recalls. The process required thinking about what he wanted, planning how he would get it and then implementing the plan and collecting data. This was followed by an evaluation of the data and revisions to the plan based on the results. This process defines the field of applied animal psychology that the Brelands had created based on Skinner’s work. It’s something that most dog trainers are ill-equipped to do.

Whenever Prins got stuck, he fell back on his old habit of blaming the dog instead of recognizing that he had signaled the wrong behavior with his body language or had poor timing or an inadequate shaping plan. According to Bob, the traditional method of training would advise, “Get a bigger stick and beat the dog harder.” He reminded Prins that he needed to stop blaming the dog and look more carefully at video evidence to see what was going wrong.

“He had been training under the eye of other trainers, who for many years [had taught] him it was the dog’s fault, and you must correct the dog,” says Bob. “If you’re the one making the errors, you should be beating yourself, not the dog.”

The three-year process was a challenge, but he kept at it because he felt that it was the only way they could get the consistency and reliability they needed. As Prins explains, “If you have a punishment-trained dog, in the new situation when they are not sure what to do, they are afraid they will receive punishment, even if it is mild. Dogs just stop performing, [and] learning slows down or stops.” He had already found that it was much more effective to condition an animal to see the world as an environment in which something positive could occur at any moment.

So he stuck with it until he had the techniques down. As a result, the program was even more successful than anticipated. “Our dogs often work far from our position, often in the dark and always in an area they have never seen before,” he says. While trainers prepare the dogs for many situations, they can never truly simulate real-time operations, which usually happen in unpredictable surroundings and are stressful for the human handlers. But once they are taught by teaching dogs that performing in many different situations is fun, dogs are able to perform reliably.

Training speed has also improved. “[With] the first dog, [it] took me eight months to train him to follow a laser. With operant-conditioning, it now takes me four weeks.”

The training is heavily weighted toward positive reinforcement, but both Bailey and Prins point out that rarely, aversives are also used. Aversives are not used until trainers understand operant conditioning well and have been training extensively in it for six months, and only when a dog exhibits behavior that puts himself, humans or the operation at risk. The aversive may range from verbal reprimands to low-level shock. Before trainers use an electronic collar, they must wear the collar around their own necks and see what it’s like to be trained this way. They find out what it feels like when a correction is given, and even worse, given at the wrong moment as commonly happens even with the most skilled trainers. “Then they understand how difficult it is, and they do not like to use it,” says Prins. Overall, aversive methods comprise about 1/1000 of the training.

Their success has led to other countries, including Belgium and Norway, adopting this approach. Despite the advantage of being able to learn from Prins’s mistakes, all the trainers in his group experience some of what he did during his first three years. To select new trainers, he sends potential candidates through four five-day chicken training camps in Sweden. “The punishment trainers fall hard. We give them four days to see if they can make the change. The process is grueling,” he observes.

The change is worth it. Trainers see the difference, and the proof is not just in their impressions. It’s in the hard data: shorter training times, more dogs trained for less money, behaviors they could never train before and more consistent, reliable dogs, which lead to more successful missions.

Bailey explains that while a handful of trainers may be motivated to improve the treatment of their dog, “the trainers who actually make the changes are those who want more success and recognize that simply beating their dog more will not get that additional success.” Once they understand and become skilled at the operant technology, an added benefit is that they can finally enjoy their work and so can the dog.

Good Dog: Studies & Research
Dog Smart: Exploring the Canine Mind
Researchers exploring the canine point of view
Canine Mind

Doesn’t it sound reasonable to study the behavior of cranes? After all, cranes are quite different from humans — they can f ly, spend lots of time on one leg and don’t need an external GPS to find their way to Florida.

But what if we replace “cranes” with “dogs”? Why study dog behavior? Unlike cranes, dogs are not a wild species with feathers, migratory patterns or conservation needs. Dogs have lived alongside humans for at least 15,000 years; are ubiquitous in human cultures; and regularly find their way into our literature, hearts and beds (also unassisted by GPS, it must be noted). We think we know dogs just fine. What’s the point of all this scrutiny?

Dogs aren’t new in the world of research.
Our shared mammalian physiology has given us reasons to open up dogs for inspection (literally) at least since the time of Descartes. In the previous century, dogs were the star subjects of Pavlov’s work on the “conditioned ref lex” (you remember: bell + food (then repeat) = dog salivates in anticipation of food after hearing bell).

In other academic arenas, Marc Bekoff, Ian Dunbar, Michael Fox and the late Frank Beach all conducted extensive investigations into canine social behavior, physiology and development. And of course, in 1965, Scott and Fuller produced their seminal text, Genetics and the Social Behavior of the Dog. Konrad Lorenz, Nobel Prize–winner and acclaimed ethologist, also had dogs on the brain; if you doubt that, pick up a copy of his book, Man Meets Dog.

Despite the thousands of years dogs and humans had spent in close proximity, scientists had never explored either the relationship or the factors that allowed dogs to become our social partners.

Dogs’ perspective as members of the human environment was missing from the equation. “It’s odd that this companion animal who has been at our side longer than any other is really not well understood,” observes Kristina Pattison, researcher at the University of Kentucky’s Comparative Cognition Laboratory.

“Dogs suffer from a failure of imagination by those asking the questions,” explains Mary Lee Nitschke, professor of psychology, Certified Applied Animal Behaviorist (CAAB) and founding member of the Association of Pet Dog Trainers (APDT). “If you already ‘know’ a dog can’t think, you’re not going to ask whether it can think.”

In 1994, imagination and an open mind prompted the creation of the Family Dog Project at Eötvös Loránd University in Budapest, initially under the guidance of Vilmos Csányi, and now headed by Ádám Miklósi. Rather than taking the position that dogs’ place among humans was unworthy of scientific investigation, they put the dog, and the dog-human relationship, under the microscope.

While previous studies had investigated owners’ intimate feelings toward dogs, the dog’s perspective on this relationship had not received comparable attention (probably because canine penmanship is quite poor and they rarely complete questionnaires in a timely fashion).

To explore the dog’s perspective, the Budapest group placed companion dogs and their owners in a modified version of the Strange Situation Test, a behavioral experiment initially created to explore the mother-infant relationship from the infant’s perspective. The test is simple enough. In a novel environment, a dog experiences separations from and reunions with an owner and a stranger while a researcher records the dog’s behavioral changes. It turned out that dogs behaved much like human infants. The conclusion? The dogowner relationship, like the motherinfant relationship, fulfills the criteria for attachment.

This research sparked a shift in perspective and demonstrated that there’s a lot we don’t know about dogs and their relationships with humans. For example, it was commonly assumed that in order for a dog to develop an attachment bond with an owner, the dog needed to be acquired as a puppy, and within a narrow age window at that. But when the Strange Situation Test was applied to adult shelter dogs who had low or restricted human contact, these dogs also displayed attachment behavior toward designated “owners.” An additional study confirmed this; guide dogs bond with their blind owners even though their relationship forms later in the dog’s life. The takeaway message was twofold: older dogs can bond with new owners, and our assumptions about dogs are not always spot on.

Since then, scientific inquiry into dogs and the dog-human relationship has exploded. “It’s almost like dogs have been rediscovered by scientists, and there are so many different aspects they can study,” notes Mychelle Blake, APDT executive director.

Dogs have now attracted the attention of a range of disciplines, from animal behavior and psychology to evolution, genetics and veterinary behavior.

Nowadays, companion dogs, working dogs, village dogs and even shelter dogs have become study subjects.

While some are interested in contributing to a growing portfolio of insights into “the dog,” others have something else in mind. “Some of the present research does not directly pertain to dogs and the humans they live with,” notes Márta Gácsi, researcher with the Family Dog Project. “Comparative studies may be examining the roots of different human social behaviors, and not necessarily concerned with the dogs themselves.”

The Duke Canine Cognit ion Center, which is part of the university’s Evolutionary Anthropology Department, takes this multifaceted approach. Their website explains, “Our goal is to understand the flexibility and limitations of dog cognition. In doing so, we gain a window into the mind of animals as well as the evolution of our own species.”

“Research also has direct application for dogs and owners,” reminds Gácsi. Jennifer Bentlage, who’s working toward a master’s degree in the cognitive biology program at the University of Vienna’s Clever Dog Lab, agrees. “If I can explain the purpose of my research to my parents, then it’s worthwhile.” Bentlage, who is currently exploring social learning, has recruited her own dogs, Monty and Michel — shelter dogs from Spain and Greece, respectively —as test subjects for her project.

“I am very interested in the dog’s cognitive abilities because this relates very strongly to the pet owner,” explains Ian Dunbar, founder of the Association of Pet Dog Trainers (APDT), veterinarian, animal behaviorist and director of the Center for Applied Animal Behavior in Berkeley, Calif. “The importance of cognition is so that people realize who dogs are and what they can and cannot easily grasp.”

Dog cognition, a seemingly technical phrase, is simply a catch-all-term that describes dogs’ attention, memory, perception, problem-solving and mental imagery skills. As you might imagine, the questions are endless.

Crystal Thompson, a self-proclaimed dog-seminar junkie from St. Paul, Minn., thinks dog cognition research is paramount. “We have learned so much in the last few years that it behooves anyone interacting with dogs — owners, trainers, shelter workers, vet staff — to do a self-audit, to check their assumptions about dogs against what research is finding.”

We expect dogs to act a certain way, and they do.
Pet owners do not expect rabbits or cats to act as social partners, but we do expect that of dogs — and rightfully so. “Through selective pressures, we pushed dogs very quickly to be highly social, and they tend to live up to our social demands,” explains Pattison.

Dogs have the potential to move in tandem with humans, stopping when we stop and starting again at our first subtle sign of forward motion, but magic is not the mechanism (although the resulting experience can certainly feel magical!). Instead, research finds that dogs are astute surveyors of human behavior, and everything from our gaze to our larger body movements provides meaningful information.

For example, researchers have found that the right side of the human face better expresses our emotional state, and when looking at other humans, we display what’s called a left-gaze bias, or a propensity to look toward the right side of the other person’s face, where all the clues are. When dogs look at human faces, they also display a left-gaze bias. Could your dog be sensitive to your emotional state? Yup.

And just as social contexts and relationships matter to you, they matter to your dog. Research has shown that if a stranger approaches your dog in a threatening manner, your dog will respond with avoidance/aggressive behavior. But if you approach your dog in the same way, your dog responds with tolerance and contact-seeking behavior. In short, dog owners feel connected to and supported by the dogs they live with, and for good reason.

But sometimes, dogs don’t act the way we want them to.
When dogs are out of sync with us, we wonder why. “We bring dogs into our households and expect them to understand and behave like us primates … well, primate plus plush toy,” laughs Pattison.

This is where cognition research comes in. Is the dog’s assessment of a situation comparable to the human’s, and if not, what is the dog’s vantage point?

Numerous studies have found that dogs attend to human communicative intent. As your significant other may have told you (possibly more than once), how you say it matters. Want a dog to come to you from across the room? Research by Patricia B. McConnell, and beloved Bark columnist, explored how different sounds affect dog behavior: “Four short notes were more effective at eliciting a come response and increasing motor activity levels than one longer continuous note.” In the real world, yelling “COOOME!” (akin to one longer continuous note) will most likely result in exasperation, but short, rapidly repeating notes, like “Pup-pup-pup,” will likely bring a dog on over.

Are dogs cooperative or competitive? Are dogs like chimpanzees, who more readily locate food in competitive situations when prohibited from going to a certain location. (A possible chimp translation might be: “The only reason you’re telling me not to go there is because that’s where the good stuff is, you jerk.”) Nope, dogs fare better in cooperative situations, finding food when informed of its location in a friendly, cooperative tone. Bringing this back to daily life: if you wonder whether the dog-human relationship is based on competition or cooperation, this is another check mark on the cooperative ballot.

Understanding the dog’s perspective is important because incorrect assessments of behavior can cause problems in dog-human relationships. “People think the dog is doing something to create trouble,” explains Floridabased Soraya Juarbe-Diaz, DVM, and board-certified veterinary behaviorist. “Almost everything on TV is about conf lict — a fight, someone trying to win or cheat; it’s very confrontational and we are always looking for a fight for superiority. Fortunately, animals don’t always work that way.”

And why can’t dogs just do what we want them to do? In one study, dogs who performed a 10-minute sit-stay (meaning, they exerted self-control) gave up quicker on a subsequent task than dogs not required to exert that initial self-control. This initial act of controlling their behavior depleted their energy, thereby adversely affecting subsequent behavior.

The consequences for not understanding dogs’ behavioral capacities can be dire. “The biggest cause of death in dogs is behavior problems, and failure to inhibit is at the root of many behavior problems,” explains Pattison.

“He barks all the time — failure to inhibit barking; he growled — failure to inhibit an aggressive response. He jumps up — failure to inhibit jumping. We expect dogs to forgo their species-typical behavior patterns and inhibit them in favor of a response we find more appropriate.” The applications for dog cognition research are vast. “The cognitive research says, look, the dog is not doing this to get your goat, he’s not doing this to diss you, he’s not trying to dominate you and take over the world,” says Dunbar. In other words, he’s just being a dog. Kelly Ballantyne, DVM, finds that the “final showdown” misconception often results in inappropriate owner-dog interactions and worsening of the dog’s behavior.

Dog-care professionals are taking note of this growing body of research. In 2004, APDT began including a science track at their yearly conference. Their mission is not simply to saturate trainers with research, but also to provide education about research methods and the nuances of study design. APDT leadership hopes that as more research hits the press, trainers will read studies with a critical eye and make informed decisions about their application for training. As Blake points out, “One study simply advances a hypothesis, and ongoing research is needed to improve our understanding of dogs.” “Research helps broaden our view of what is possible,” add Nitschke. “The fact that cognitive studies sometimes disagree with each other is wonderful, because it means there is more for us to look at. Different studies with different results broaden our vision of what is possible.”

Research also has application in the classroom. “My students were blown away,” reports Johnna Chamberlain, who teaches at the Lang School in New York City, a school for gifted children with learning differences. That’s how she describes her students’ reaction to “Dogs Decoded,” the NOVA special on dog cognition. “[My students] hadn’t considered that dogs were communicating through their barks, bodies or tails. Since [my students] might have social or emotional delays in relation to perspective taking, it was a big deal for them to consider their own relationships with pets and realize that dogs are communicating and thinking in their own unique way.”

Thompson looks at the breadth of cognition research from a different angle; she wonders whether owners are providing for their dogs’ mental needs. “It’s important for owners to realize that dogs have real mental abilities and needs. Putting food in a dog’s bowl is just wasting his brain. It’s the little things — Kongs, Tug-a-Jugs, hiding kibble around the house — it’s not hard, and it’s a simple way to engage their natural abilities.”

“How you interact with dogs often depends on your impression of their abilities,” notes Juarbe-Diaz. Reframing our expectations, with a little help from research, could set up dogs to succeed in our primate world. As Gertrude Stein said, “I am because my little dog knows me.” Shouldn’t we do the same for dogs?

Good Dog: Studies & Research
Q&A with Mark Derr about Dog Origins
Venturing deep into the history of our oldest friends
sociable wolves

Mark derr, long-time Bark contributor and historian of the dog, recently released a new book, How the Dog Became the Dog, in which he examines canine evolution. Derr covers a lot of ground in this work — 135,000 years, to be precise! We talked with him about the dawn of dog, and how our evolutionary pathway coincided with theirs.

Claudia Kawczynska: Canines going from fierce predator to “loyal companion” is quite a leap. Can you sketch how and why this might have happened?
Mark Derr: The premise I start with is that, in many ways, dogs are an evolutionary inevitability. As soon as humans and wolves encountered one another on the game trails, they struck up a relationship, and they’ve been at it ever since. People and dogs have very similar social structures, and there was a level of sociability between some wolves and some people that allowed those particular individuals to come together.

So, wolves and humans had an affinity, and sociable wolves would often breed near human societies. As they began to do that, populations were established, though not everywhere and not in great numbers. One group of socialized wolves would die out and others would appear in other places at other times. There is evidence that destroying the structure of a wolf pack destroys the culture for the young, leaving them without guidance. Imagine that this happened over many, many generations, resulting in a more socialized “dogwolf” — or dog-like wolf. In that sense, you’re never going to find a single place for the [first domesticated] dog to have appeared. Rather, you have [the dog developing] wherever you have wolves and humans.

CK: What was in it for the wolves who paired up with us?
MD: Several things. Wolves hanging around humans probably ate better than “wild” wolves. They also gained sexual freedom, which is a positive thing. Raising puppies takes a lot of time and energy, but with humans around, the burden of raising the young was greatly diminished.

CK: Was this consciously directed?
MD: No, not on the part of the wolf or the human, who didn’t think about it as a benefit either. The operational presupposition is that animals don’t do things that are bad for them. So if you look at it, let’s face it … if someone comes along to help raise your offspring, you are suddenly free to do all kinds of stuff.

CK: Like getting better food to feed those offspring.
MD: I think that was a major benefit for the dogwolf. Besides, they received protection from us.

CK: Did this relationship affect our own evolution?
MD: There is no question that early humans benefited from dogs at a very fundamental level. Evidence shows that humans who hunt with dogs do much better than those who don’t, for example. So, theoretically, the quality of their food would be better. The dog also provided transport, greatly extending our reach, and protection. Having that barking wolf or dogwolf or dog alert us to strangers, who may not have had our best interests at heart, was a positive thing — it helped us stay alive to reproduce. There is also no question that dogs helped us hunt, and then control and guard, ungulates that had been domesticated by humans. Dogs have been invaluable to humans, and continue to be.

CK: You note that with genetic data pushing back the dawn of dog to perhaps 135,000 years ago, the idea of neoteny has been turned on its head. How so?
MD: Though dogs reach sexual maturity earlier than wolves, growth or development of other organs and limbs is delayed, which is called paedomorphism. Slowing the rate of development is said to lead to neoteny, the retention of juvenile traits into adulthood.

Now, the latest research shows that a small number of genes have a big effect on everything from overall size and leg length to numerous other factors. I’ve been saying that for years. I didn’t know precisely what the mechanism was, but it wasn’t paedomorphism. There are other explanations, as it turns out. Not only that, but many of the features some consider neotenic are simply creations of modern breeders, who strove to make dogs more cuddly and humanlike by selecting for rounded skulls and large, forward-facing eyes.

To reiterate an important point, which I’ve made in numerous articles in The Bark and in my books, there is no evidence that dogs originated from selftamed, submissive, neotenic wolves. That theory — which is based on dogs originating during the Mesolithic Age, when people lived in settlements with garbage dumps — is not right. Dogs evolved much earlier than that and were in the camps with the hunters and gatherers.

CK: Back when we were both hunting the same species — together or separately — is it possible that wolves were hunting us too?
MD: I don’t know of any record of wolves hunting humans, though humans have long hunted wolves. The Plains Indians had wolves around the bison they were hunting, and there is strong evidence that shows their dogs regularly crossbred with the wolves. And sometimes to the point where it was not possible to distinguish between wolf and dog. The thing about human behavior is that much of it is fixed; even the tools we use are basically the same. It might be made from different materials, but a knife is still a knife.

Take an example from Lewis and Clark, who describe great herds of bison out there on the plains, and the Indians who hunted them with their dogs. The wolf is described by Lewis as the “shepherd” of the bison. The way wolves hunt really isn’t that much different than the way herding dogs gather animals.

People like Barry Lopez have done work on the business of wolves and human cultures, and why wolves are so distrusted by humans. The answer, I suspect, is that once dogs and agriculture were firmly established, a divorce occurred between humans and wolves (and other wildlife), because those animals were seen as threatening our livelihoods. At some point, the wolf became a competitor — an enemy, even — not because it was hunting us, but because it was taking our livestock. The mediating force is the dog.

More recently, the conservation movement established a sharp divide between the wild and the built, a divide that really shouldn’t exist, but does. At that point, the wolf became one thing and the dog became another, and they are in opposition rather than what they are, which is very closely related. I don’t think that the wolf has ever been an enemy of humans, but I could be wrong.

CK: Pat Shipman, archaeologist and author of The Animal Connection, pointed out that it was unlikely that wolf packs tracked nomadic hunters in order to live off their spoils (among other things) because those wolves would have had to cross the territory of other wolves, which would have been highly dangerous for them. So it was more likely that the dogwolf and the human were partners in the hunt. What’s your take on that?
MD: I agree. The other thing is — and this goes back to the food issue — humans are much more prof ligate hunters than wolves, and also take reproducing-age adults. Those are usually the strongest animals, and provide the best and most food. Wolves take the old and the young. Interestingly enough, once humans domesticated animals, we also tended to cull the very young and the beyond-reproduction-age females. So we came to eat more like wolves. I also firmly believe that humans and wolves just liked each other.

CK: Besides our shared characteristics, what else do we have in common with dogs?
MD: We are both extremely defensive of our territory, protecting home and hearth no matter where that is. We also both like to be on the move — we like going walkabout. Anybody who has dogs knows they love to come out with you. Dogs like to go for rides. Or, open a gate and the dog is gone. We value one another’s companionship, and we keep each other warm at night.

Dog's Life: Lifestyle
Humanity's Best Friend
We may owe our evolutionary success to the domestication of dogs

Scientists have long debated why Neanderthals lived successfully on earth for over 200,000 years and then died off after humans came along, about 40,000 years ago. Some blame it on climate change, while others think humans were more successful at getting food due to social cohesion and the ability to develop advanced tools.

However, a new theory claims that we may owe our evolutionary success to the domestication of dogs.

Anthropologist Pat Shipman speculates that the relationship between humans and canines began in hunting, where dogs would help people identify prey and haul carcasses back home on their backs (dogs of the Paleolithic era were typically the size of a modern-day German Shepherd or larger). This assistance gave humans an advantage in obtaining food and conserving energy.

Shipman believes that the relationship between humans and dogs only got stronger over time and may have even led to shared characteristics for communication. In humans, the whites in our eyes are highly visible, compared to other primates, allowing people to "talk" silently when hunting in groups. It turns out that canines, unlike other animals, have the ability to decipher our eye contact. In a study conducted at Central European University, dogs performed as well as human infants at following the gaze of a person when they held their head still.  

Because canines may play an important role in our success over Neanderthals, Shipman calls them "humanity's best friend." Dog lovers need no validation that our pets are critical in our lives, but now we may have more to thank them for than we previously thought!

Culture: Science & History
DNA Testing
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.

Want to read more about dogs and DNA? Click here.

Culture: Science & History
Buffon
The Science and Art of Georges-Louis Leclerc, Comte de Buffon

In setting out to explain everything that was known about the natural world in his 44-volume Histoire Naturelle, Georges-Louis Leclerc, comte de Buffon (1707–1788) reserved a special place for the dog. In fact, he may have been the first naturalist to raise the question of the role of dogs in human evolution: “To conceive the importance of this species in the order of Nature, let us suppose that it never existed. Without the assistance of the dog, how could man have conquered [and] tamed…the other animals. … The training of the dog seems to have been the first art invented by man; and the result of this art was the conquest and peaceable possession of the earth.”

Born in 1707 in Montbard, Burgundy, the son of landed gentry, Buffon was interested in many disciplines; he studied law and mathematics, and was also fascinated with the biological sciences. The first few volumes of Buffon’s groundbreaking work were published in 1749, in the middle of the French Enlightenment, a period considered to “have given birth to the modern world.” Yet it was also a time steeped in church doctrine that provided only dogmatic biblical answers to most questions about biological diversity.

His Histoire Naturelle was an ambitious and tremendously popular project; it was the most widely collected work of its time and reached more readers than even the classics of Voltaire and Rousseau. Replete with Buffon’s insightful theories on everything from the formation of Earth and the planets to crustaceans and rhinoceros, it contained beautifully executed and detailed illustrations (including those of dogs, as shown here). Arguing for a more dynamic and inclusive natural history, Buffon resisted the detailed taxonomic classifications of Linnaeus.

In his foreword to Jacques Roger’s biography, Buffon (1997), L. Pearce Williams writes that “it is no exaggeration to suggest that Buffon’s ideas stimulated much of the scientific work in natural history that culminated in the Darwinian synthesis. Even when he was wrong, as he often was, he was precise enough in his errors to permit others to see beyond his vision.”

In considering the dog, Buffon was fascinated by the many different types of canines. He recognized 30 “fixed varieties,” but acknowledged that there could be even more. He also described 17 “races”—as he called the differences within species—that he attributed to geographic influences such as climate. Into one of these groups he put the “shepherd dog, the Pomeranian, the Siberian, the Lapland and Iceland”—dogs that have “an instinct which induces them to follow and protect flocks”; into another group were “the grey-hound [sic], the large Danish dog and the Irish grey-hound.” Dogs in this latter grouping “are fond of running, and of following horses and carriages … they hunt rather by eye than the nose.”

The next grouping belonged to the “true hunting dogs,” the “hounds, harriers, spaniels, terriers and water-dogs.” Lastly, he grouped the “small Danish dog and the Turkish dog” together, and placed the British “bull-dog” in its own “group,” noting that it was difficult to “preserve” that particular breed in France—he remarked that these dogs “often send forth a disagreeable smell.” In his construction of a canine genealogical tree, he placed the “shepherd dog” at its root. He tried to mate a dog with a wolf, and a dog with a fox, but because his limited experiments were unsuccessful, he concluded that they all were distinct species and that the “dog derives not his origins from the wolf or fox.”

It was with his theory of “degeneration” of species due to climate, and most especially to the climate of the New World, that he had the most long-standing influence outside of the scientific field. In fact, modern pundits have ascribed the fractious Franco-American political relationship to Buffon’s “barkless American dog” theory. Although it was Dutch philosopher Cornelius de Pauw who first claimed that “dogs cease to bark” when brought to America, Buffon picked up on this observation and used it to partially explain the differences in Old and New World species. He thought that the latter were of the “degenerative” variety—weaker, smaller and generally more puny—hence, dogs unable to bark.

Great American thinkers, including Hamilton, Jefferson and Franklin, took umbrage at this, believing that it was meant to extend beyond “natural history” to politics—that this “American degeneracy” was also being applied to their newly formed style of government. In number 11 of the Federalist Papers, Hamilton wrote, “Men admired as profound philosophers have, in direct terms, attributed to her inhabitants a physical superiority, and have gravely asserted that all animals, and with them the human species, degenerate in America—that even dogs cease to bark after having breathed awhile in our atmosphere.”

Jefferson wrote his book Notes on the State of Virginia (1781) to refute Buffon’s theory, and included a “careful analysis of the relative sizes of American and European animals.” Buffon recanted this theory before his death, but old myths die hard. The “barklessness” of our dogs survived well into the last century, when the poet Paul Claudel, serving as the French ambassador to the United States in the 1930s, repeated this fiction.

Though most of Buffon’s theories explaining natural phenomena were, by and large, incorrect, one of the most important contributions he made—as was noted by biographer Jacques Roger—was to “transformed the way of understanding nature.” Buffon’s courageous and innovative approach paved the way for Darwin and other revolutionary thinkers responsible for much of what we know today about the natural world.

ABOUT THE ARTIST
Jacques Eustache de Sève (1742–1788) was the principal artist commissioned by Georges Buffon for his Historie Naturelle. His lovely and lifelike illustrations were complete scenes that included classical landscape backgrounds, which influenced later natural history illustrators. One writer comments, “The illustrations to Buffon’s original edition are exquisitely engraved and exude aristocratic elegance and charm. Whether the animal depicted is a lion or a pig, it is shown as if in a tableau revealed to our curious gaze by the drawing aside of a curtain. The animals do not seem like wild beasts roaming free in their native woods, deserts and mountains, but like actors performing among stage props and painted scenery for the benefit of the lords of creation.” These illustrations were engraved on copper plates and printed on paper made of cotton or linen rag, which contributes to their long-lasting qualities. The original engravings were produced in black ink, and hand-painted with watercolors centuries later.

 

Culture: Science & History
Can DNA Decipher the Mix?
Unraveling the genetic tapestry provides clues to breeds and their mixes

A mongrel dog is like a box of chocolates: You never know what you’re going to get. And therein lies the appeal. What’s more fun than serendipitous unpredictability all bundled up in puppy fur? But when that puppy grows up, we inevitably make assumptions about her ancestry based on how she looks and behaves. Our logic goes like this: “If my pooch is long and low to the ground, and she never barks, she must be a Corgi/Basenji mix.”

But it’s much more complicated. The genes—and there may be hundreds—that work together to make a Corgi look like a short-legged Shepherd may be completely different than those responsible for a Basset Hound’s low-slung carriage. With some exceptions, scientists cannot yet connect genetic dots to specific traits. But they have discovered something tangible that measures some of the differences between breeds: genetic patterns of organization displayed on a scatter graph that answer the question, “What’s the same and what’s different?”

A scatter graph provides a symbolic visualization of DNA, wherein each individual dog contributes one point. The resulting pattern indicates the type and strength of the relationship between individuals. The more the points cluster around each other, the more alike they are.

 

Breakthrough
Until only a few years ago, scientists couldn’t identify the differences in genetic material that might explain profound variations in the Canidae clan. From wolf to West Highland White Terrier—they all looked the same under the microscope. Then, in 2004, Elaine Ostrander and her colleagues at the Washington-based Fred Hutchinson Cancer Research Center published data indicating that as much as 30 percent of the dog’s genetic material accounts for breed variation (Science, May 2004).

In addition to simplifying methods used to find markers for breed-related disease, the researchers identified patterns of “breedness” and tracked the history of breed DNA. At the same time, by following mitochondrial DNA, genetic material passed down from mother to offspring without changing, they traced the breed’s journey.

Depending on how much time is attributed to a generation and how many generations are involved, scientists can estimate how much time has passed. Based on this tracking, it has been suggested that it took 5,000 years to develop and refine a handful of the world’s 350-plus breeds, and about 400 years to create the rest.

Research indicates that four distinct breed groups are ancient: (1) Middle Eastern Saluki and Afghan, (2) Tibetan Terrier and Lhasa Apso, (3) Chinese Chow Chow, Pekingese, Shar-Pei and Shih Tzu; Japanese Akita and Shiba Inu, (4) Arctic Alaskan Malamute, Siberian Husky and Samoyed. Although the 13 breeds look different, they are so closely related that they are represented by a single genetic cluster. It’s likely they all originated from the same stem-parent—proto-breed, if you will—who roamed the Asian continent.

As humans migrated from one place to another, this ubiquitous proto-breed trotted along, bringing with her the ingredients needed to cook up all the breeds we’re familiar with today. Her offspring performed work unique to each geographical region, such as hunting, hauling or guarding. Isolated and mating only with each other, “accidental” breed types exhibiting consistent shape, color and behavior emerged.

No matter what historians might claim—scent hound to sight hound, bird dog to bad dog—evidence produced through genetic research indicates that all remaining breeds have been concocted in the last 400 years. Although closely related to one another, they can be identified as distinct based on the way their DNA separates.

How They Do It
Sue DeNise, vice president of genomic research at MMI Genomics Inc., which developed the Canine Heritage Breed Test specifically for mixed-breed analysis, talked to us about how her company analyzes canine DNA. “We’ve been doing testing for AKC parentage verification for a long time,” she notes. “We initially started working in the cattle business, looking for genetic markers in order to trace what was important to cattle breeders. Out of that whole-genome association study, we had purebred and crossbred cattle, so we asked, ‘What can the markers tell us about underlying traits in breeds of cattle?’” Their discovery paved the way for the companion animal program, which was modeled on what they learned with cattle.

“We look at ’breedness‘ among dogs. Our canine database is built with 10,000 samples of 108 breeds. We ran 400 markers to identify the best markers for a ’breedness‘ test against 38 breeds. We created a panel of 96 pieces of DNA that split dogs into their identified pure breed. In our preliminary test, we found that individual purebred dogs cluster with other purebreds.” Initially, MMI chose 38 AKC registered breeds from their database, selected for their popularity based on number of registrations. Recently, as DeNise notes, they increased the number to 108 breeds. This jump in breed recognition required testing thousands of markers to identify the just over 300 markers that characterize these 108 breeds.

Constructing Breeds
Like all species, domestic dogs are on an evolutionary journey, starting at wolf and going somewhere yet to be determined. We tinker with evolution, but might be surprised to find out we don’t control it. Our concept of a breed—that individuals within the breed look alike—is nothing more than a snapshot of the DNA time line, taken while we’re doing the tinkering.

Breeds are created a number of ways. In simple terms, when breeders interfere with natural reproduction and rigorously select for traits favored by humans, specialized breeds like Retrievers, Spaniels, Hounds and Terriers are the result. Saving spontaneous mutations in a litter of dogs, repeating the breeding to get more of the same mutation, and breeding those dogs back to one another has resulted in the English Bulldog, Chinese Crested and Inca Hairless. More recent breeds, such as the Airedale, Australian Cattle Dog and Doberman, are the result of crossing older breeds to make new ones.

When kennel clubs closed gene pools in the late 19th century to suspend change in registered dogs, breeds drifted toward a more uniform stereotype. Until the early 1800s, an assortment of dogs with similar talents who could produce somewhat similar offspring were awarded the right be called a breed. Breeds evolved, flourished and disappeared when jobs were eliminated. Tumblers, who mesmerized prey by “winding their bodies about circularly, and then fiercely and violently venturing on the beast,” disappeared when guns came into widespread use. Turnspit dogs, who made a living running on a wheel to turn meat so it would cook evenly, received their pink slips when technology improved cooking methods.

By and large, Victorian society was not so pragmatic; sentimentality and commercial opportunity were catalysts for saving unemployed breeds from their inevitable demise. As a result, many Terrier breeds went from killing varmints in the barnyard to killing time in the Victorian parlor in less than a decade.

Whereas previously, a breed was a regional product maintained and preserved by a small community of knowledgeable people, commercial interest in the well-bred pet dog initiated a shift in breeding practices during the Victorian era. The old-money kennels operated as a pastime by the wealthy gave way to a large number of small, commercially operated kennels run by entrepreneurs of modest means and experience.

Germane to this tale is that, according to the unwritten rules governing canine physiology, anatomy and behavior go hand in hand. One cannot be changed without affecting the other. Victorian enthusiasts who were busily adding aesthetic traits to utilitarian breeds were creating not only subtle variations in type, but in many cases, modifications in behavior as well. As utilitarian breeds went from working hard to hardly working, many exhibited new physical and behavioral characteristics that were compatible with their augmented duties as companion animals. Breeders claimed the “sub-breeds” as their own, made up new names and registered each one.

However, no matter how they’re sliced and diced, reducing and suppressing genes so they aren’t expressed doesn’t mean they’ve been eliminated. They’re still lurking and, depending on the method used to analyze the DNA, the lurkers often show up in the results.

Deconstructing Breeds
The problematic aspect of analyzing mongrel DNA is that breeds were not all created at the same time. As DeNise explains, “As new breeds are developed, they may not appear as uniform as older breeds. When older breeds are crossed to create a new breed, there is some period of time before the new breed develops a unique DNA pattern of its own. In these cases, the more ancient breed sometimes appears in the new breed. The number of generations required to have a uniquely identified breed created from crossing of older breeds depends on the number of breeding animals in the new line, the severity with which the breed owners apply the standard, and the amount of introgressing [inbreeding, or breeding immediate relatives; line breeding, breeding close relatives; and backcrossing, breeding sibling to parent] allowed by the registration agency.”

Most people assume all mixed-breed dogs had a purebred ancestor at some time in their recent heritage. But in fact, this is not necessarily the case. When you run a mongrel’s DNA through a computer program, the algorithms attempt to group breeds together on a scatter chart. If the heritage of the dog is such that it is not in MMI’s database of 108 breeds, the program tries to find varieties that are most alike. Because at least one or two of the handful of ancient breeds are in every modern dog, sometimes the program will identify an ancient breed in the mix. “In the report we send to the client, we use the terms ‘primary,’ meaning the majority of the DNA matches a breed; ‘secondary,’ meaning less than the majority of the DNA but a strong influence nonetheless; and ‘in the mix,’ meaning the least amount of influence,” DeNise notes. That’s how you might get an obscure breed in the report. For instance, a 35-pound mongrel with a tablespoon of Husky and a teaspoon of Border Collie may also have a dash of Borzoi, because before gene pools were closed a century ago, Huskies were crossed with coursing hounds to add speed.

Don’t Judge a Pup by Her Cover
As MMI Genomics states on their certificates, “Your dog’s visual appearance may vary from the listed breed(s) due to the inherent randomness of phenotypic expression in every individual.” What this means is that you may look nothing like your parents, but you have Grandma’s great legs and Great Uncle Harry’s turned-up nose. All in all, though, no matter how genes are mixed and matched, your family members resemble one another. However, if Grandma was an Afghan Hound and Great Uncle Harry was a Pug, “random phenotypic expression” can be pretty extreme.

Researchers are intrigued by data that suggest expressed traits are somehow turned “on” and “off” by other genetic components, thus causing the wide variations in canine form and behavior. For instance, it’s possible that many breeds have the genetic potential for a black tongue, but only a few breeds have the molecular mechanism to switch that color on. So that black-tongued mutt may not have any Chow in the mix after all.

On the other hand, the results may show that a quintessential Heinz 57 has the genetic makeup of a single breed and it could be one she looks nothing like.

DeNise explains it this way: “In a population of any breed, there are dogs that are carriers but don’t exhibit phenotype [observable characteristics]. If you reduce the size of breeding population—creating what we call a bottleneck—you start increasing the frequency of deleterious traits, like dwarfism or white coat. If we looked at the DNA of, for instance, a group of white mini-German Shepherds, they would probably cluster with German Shepherds. After they’ve intermated for five to six generations, we may not come up with that. They would cluster with each other. If breeders were changing allele frequencies quickly, you could do it very fast.

“There are always contradictions that make you say, ‘Huh, that’s really weird.’ One odd thing that happens is due to some sort of random assortment of genes in mixed-breed dogs. The algorithm may identify a breed that is not consistent with the physical appearance of the dog. We sometimes get an indication of this when the certificate is printed with the picture of the dog provided by the owner, and the certificate is reviewed by our customer service department prior to mailing it to the pet owner.”

A 90-pound, wiry-haired mongrel who swims, chases balls and makes goo-goo eyes like a Golden Retriever and whose only pedigreed relative is a very distant Chihuahua confounds the process, but says a whole lot about the complexity of canine genetics and why some scientists devote their careers to studying canine evolution. Extreme variation in anatomy and behavior is unique to the domestic dog. If humans were an equally anomalous species, we’d weigh between 20 and 650 pounds and range in height from three to 10 feet. In dogs, adaptations change with such speed that scientists suspect there may be a clue in the canine genome that could reveal how evolution works.

Sorting It Out
Before they launched the project, MMI tested DNA from a street dog rescued from a Thai village. You’d think there would be no clusters of any kind, but the computer identified Chow and Akita in the mix. This isn’t surprising, because the free-living common village cur who populates most of the developing world may be the closest living relative to the original proto-breed. Findings suggest that Thai pooch stores a sizable chunk of the original genetic blueprint of every single living dog in her DNA. The question is, how much?

MMI can’t as yet define the percentage of “breedness” in mixed-breed dogs. One reason is that some breeds cluster loose and others tight. Why this happens isn’t clearly understood. German Shepherds, Standard Poodles and Collies cluster tight. Miniature Poodles cluster tight, but Toy Poodles cluster loose. Within their breeds, Labrador Retrievers and Beagles often cluster as two different groups. According to DeNise, “Labs from the United Kennel Club that are bred specifically for hunting and AKC Labs do not necessarily cluster as one breed. And AKC Beagles and Beagles bred specifically for research don’t cluster together either.”

She adds, “In my opinion, it’s possible that a population that increases rapidly doesn’t cluster as well as those populations that have remained static. This is because, as you increase a population to accommodate breed popularity, people breed everything, including animals that may not exhibit all the physical characteristics that are desirable.”

And even when dogs look alike, they can display behavioral differences. As DeNise notes, “We understand so little about how behaviors are coded. Many behaviors are learned, but there are probably multiple genes that are responsible for herding, birding, heeling—these kind of hard-wired behaviors.”

Scientists are eager to tease out genetic connections to breed-associated motor patterns. When wolves hunt, they display these behaviors sequentially: orient > eye > stalk > chase > grab-bite > kill-bite > eat. Artificial selection, however, extracts and segregates these patterns in incomplete sequences. In certain breeds, individuals perform the abbreviated motor pattern repeatedly. A Pointer who stops dead in her tracks and stands stock still with her front leg held rigid in mid-stride to indicate the presence and position of game is the lofty goal of bird-dog breeders. To wolves, it’s just a good meal interrupted.

Combinations of canine anatomy and behavior push and pull one another along in a rhythm of interconnected patterns in relationships that may not be as random as they appear. Like principal components of an automobile in which the size of the engine and the weight of the body directly affect efficiency, it appears that dogs, too, have integral parts wherein one component is proportionate to the other.

Researchers don’t fully understand the relationship, but they are making headway. As reported in Genetics (June 2008), a team of scientists identified a few single genes that regulate systems controlling skull shape, weight, fur length, age span and behavior. Because mutts are combinations of DNA from different breeds, they may hold the answer to how the genes influence multiple traits.

Scientists suspect that many evolutionary secrets are hidden in the dog genome. For dog lovers, deconstructing Molly or Max’s mixed-breed heritage is an interesting intellectual mystery to be discussed at cocktail parties or the dog park. For scientists, their genetic material is nothing less than an instruction manual for species building. Whereas populations evolve over the course of millennia through the process of natural selection, dogs can change so rapidly and abruptly that they represent evolution at hyperspeed. How it happens remains a puzzle. Now scientists are looking to mutts to find the missing piece.

 

 

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