research on behavior of dogs

Dog study shows there's a lot more to behavior than just breed

When comparing dog breeds and mutts, distinctive physical traits are not highly correlated to inheritable behavior traits.

A genetic study of 2,155 purebred and mixed-breed dogs combined with 18,385 owner surveys has challenged existing notions about dog breed stereotypes and personality types. The study identified 11 locations along the canine genome that were strongly associated with behavior, none of which were specific for breed, suggesting that these personality traits predate modern canine breeding by humans.  

“Using a really powerful model, these findings provide compelling support for the fact that complex traits, like behavior or personality, may have some genetic contribution, but that contribution is distributed across so many genes with really tiny effects," said Arizona State University School of Life Sciences Assistant Professor Noah Snyder-Mackler , who contributed to the study in a collaboration with investigators at the UMass Chan Medical School, including Elinor Karlsson, who led the study that was published in a recent issue of Science. "This means that while we can predict how biddable Biddability refers to a dog's ability to respond to human direction. a dog is on average, we will do a pretty terrible job predicting the biddability of a single dog based on their genotype, and definitely not their breed. It points to the fact that much of these complex traits are strongly impacted by the lived experiences of each individual.”

Noah Snyder-Mackler

Canine behavioral disorders are often proposed as a natural model for human neuropsychiatric disorders. Compulsive disorders, for instance, are often observed to manifest similarly in both humans and dogs. Thanks to the power of current DNA sequencing technology and the close relationship between pet and owner, canine genome-wide association studies (GWAS ) have the potential to identify unique genetic areas in the dog genome that could lead to new insights into similar genes in humans. Karlsson and colleagues show that large-scale GWAS in dogs can yield genetic loci associated with behavioral traits.

“Although friendliness is the trait we commonly associate with golden retrievers, what we found is that the defining criteria of a golden retriever — what makes a golden retriever a golden retriever — are its physical characteristics, the shape of its ears, the color and quality of its fur, its size; not whether it is friendly,” said Karlsson, associate professor of molecular medicine at UMass Chan Medical School.

“While genetics plays a role in the personality of any individual dog, the specific dog breed is not a good predictor of those traits,” explained Karlsson, who is also the director of the Vertebrate Genomics Group at the Broad Institute of the Massachusetts Institute of Technology and Harvard University. “A dog’s personality and behavior are shaped by many genes as well as their life experiences. This makes them difficult traits to select for through breeding. For the most part, pure breeds are only subtly different from other dogs. A golden retriever is only marginally more likely to be more friendly than a mixed-breed or another purebred dog, such as a dachshund.”

A dog's tale

The story of how modern-day dog breeds emerged is a relatively short one in evolutionary terms, contrasted against the history of dog domestication from prehistoric wolves. Genetic research pegs the change from wolf to dog at about 10,000 to 15,000 years ago. Humans didn’t begin intentionally breeding dogs until roughly 2,000 years ago, when they were being selected for work roles such as hunting, guarding and herding. It wasn’t until the Victorian era in the 1800s that humans began selecting dogs consistently for the physical appearance and aesthetic traits that today we commonly associate with modern breeds.

Yet modern dog breeds are often credited with characteristics and temperaments (bold, affectionate, friendly, trainable) that correlate to their ancestral function (herding, guarding or hunting). Likewise, the breed ancestry of dogs is assumed to be predictive of temperament and behavior. DNA tests are even marketed as tools for dog owners to learn about a pet’s individual personality. However, there is a lack of genetic studies linking behavioral tendencies to ancestry or other genetic, heritable factors.

By pairing genome-wide association mapping technologies with pet owner surveys obtained through Darwin’s Ark, an open-source database of owner-reported canine traits and behaviors, Karlsson and first author Kathleen Morrill, a PhD student in the Morningside Graduate School of Biomedical Sciences at UMass Chan, explored the complicated relationship between modern canine breeds and behavioral characteristics.  

Genome-wide association studies (GWAS) are an approach used in genetics research to try to associate specific areas of variation in the human genome that align with certain phenotypes. Taking whole genome sequences from hundreds of thousands of people with the same disease, for instance, researchers look for common genetic variations among these people to pinpoint broad areas in the genome that may be predictive or causative for specific diseases.

Karlsson, Morrill and their colleagues applied this same strategy to correlate areas of the canine genome with certain behavioral traits or characteristics. Pet owners who participate in Darwin’s Dogs provide scientists with saliva samples from their dogs. Researchers run whole genome sequencing on these samples to generate a robust genetic data set for investigation. Additionally, for this study, owners filled out 12 short surveys totaling 117 questions about their pet’s behaviors and physical traits. Combined, this data provides the basis for scientists to associate genetics with owner-reported behavior. 

“Given a large enough sample size, GWAS are a really powerful tool for learning about genetics,” said Morrill. “We only get that size by looking at all dogs – not just purebred dogs, but mixed-breed dogs too. We compare all these DNA sequences computationally, using complex algorithms, to identify areas of differences and commonalities that might be of interest.”

Karlsson and Morrill collected more than 2,000 canine genomes and 200,000 survey answers through Darwin’s Dogs. Because of existing stereotypes about dog behavior and breeds, Karlsson and Morrill designed the study to account for possible owner bias, in part, by establishing standard definitions for reporting and rating canine behavioral traits such as biddability (a dog response to human direction), dog-human sociability (a dog’s comfort with people, including strangers) and toy-directed motor patterns (a dog’s interest and interaction with toys), among others. Physical and aesthetic trait standards were pulled from those published by the American Kennel Club.

Genetic research pegs the change from wolf to dog at about 10,000 to 15,000 years ago. Photo courtesy of Noah Snyder-Mackler

More to behavior than just breed

Behavioral data was analyzed across owner-reported breeds and genetically detected breed ancestries. The results of these tests, which included data from 78 breeds, showed that while breed explained some minor variation in behavior, its contributions were relatively small (9%). For certain behavioral traits, such as toy-directed motor patterns, age was a better predictor of behavior: Younger dogs were more likely to score higher in this category. For specific survey items, such as “lifts leg to urinate,” a dog’s sex was the best predictor of behavior. Physical traits like coat color were more than five times more likely to be predicted by breed than behavioral traits.

Additionally, investigators failed to find behaviors that were exclusive to any one breed. Even in Labrador retrievers, which had the lowest propensity for howling, 8% of owners reported their Labradors sometimes howl. Likewise, while 90% of greyhound owners reported that their dogs never bury their toys, three owners described greyhound dogs as frequent buriers.

Complementing the survey analysis of breeds to measure breed-behavioral propensities, the researchers also leveraged the genetic ancestry of highly mixed-breed dogs to test whether behavior is heritable in a breed-dependent manner. In some cases, heritable behavioral traits like biddability are somewhat more likely to correlate with breed, even if mixed a few generations back. In the case of purebred dogs, ancestry can make behavioral predictions somewhat more accurate. For less heritable, less breed-differentiated traits, like agonistics threshold, which measures how easily a dog is provoked by frightening, uncomfortable or annoying stimuli, breed is almost useless as a predictor of behavior.

A comparison of dog genomes was performed to identify genetic variations tracking along breed, as well as along individual physical and behavioral traits. Karlsson and Morrill identified 11 locations on the dog genome strongly associated with behavioral differences, none of which were specific for breed, and another 136 suggestively associated with behavior. The genetic differences between breeds such as golden retrievers, Chihuahuas, Labrador retrievers, German shepherd dogs and others, primarily affected genes that control coat color, fur length and other physical traits — far more than breed differences affected behavioral genes.

Overall, Karlsson and Morrill found that behavioral characteristics were influenced by multiple factors, including environment and individual genetics, but that modern breed classification played a modest role in the outcome.

“The majority of behaviors that we think of as characteristics of specific modern dog breeds have most likely come about from thousands of years of evolution from wolf to wild canine to domesticated dog, and finally to modern breeds,” said Karlsson. “These heritable traits predate our concept of modern dog breeds by thousands of years. Each breed inherited the genetic variation carried by those ancient dogs, but not always at exactly the same frequencies. Today, those differences show up as differences in personality and behavior seen in some, but not all, dogs from a breed.”

Next up: Understanding 'dog years' 

For ASU's Snyder-Mackler, the study represents a significant opening salvo demonstrating the power of using modern genomic tools to study companion animals to gain new insights into the causes and consequences of variation in the social environment, from the molecular to the organismal levels.

"This study would not have been possible without the Herculean efforts to generate a massive behavioral dataset and combine that with whole genome sequencing of almost 2,000 dogs – without spending an arm and a leg. This was achieved by using some really cool computational approaches to try and gain as much information from each dog with as little sequencing as possible, which we found worked better than the most common approach to genotyping — microarrays," said Snyder-Mackler. His lab focuses on studies that allow him to probe questions central to human health, aging and evolutionary biology, often using companion animals or field animal research.

Next up is another huge citizen science canine effort now underway called the Dog Aging Project, where, according to Snyder-Mackler, "We are trying to understand what makes dogs tick (and age). Our lab uses molecular tools to try and identify how age and the environment interact to alter the dog immune system. So this new study has provided a really powerful roadmap to carrying out future studies in larger cohorts, which Dr. Karlsson is leading as part of the Dog Aging Project." 

The project promises to explore what exactly are "dog years," other than the conventional wisdom of one dog year equal to about seven human years. What is known is that big dogs typically age more rapidly, an estimated 10 times faster than humans. By contrast, little dogs have a longer life span and can often live up to 20 years old. 

Given that dogs share the human environment and have a sophisticated health care system but are much shorter-lived than people, they offer a unique opportunity to identify the genetic, environmental and lifestyle factors associated with healthy life span.

Snyder-Mackler and collaborators outlined the goals of their efforts in a recent Nature paper . The Dog Aging Project is an interdisciplinary, open-data, community science project that consists of a team of more than 100 staff, students, faculty and veterinarians from more than 20 academic institutions, along with over 30,000 canine participants and their owners. 

The study investigators also include Kate E. Creevy, Joshua M. Akey, Matt Kaeberlein and Daniel E. L. Promislow, and afford great opportunities for Snyder-Mackler lab ASU graduate students, such as Brianah McCoy and Layla Brassington, who are helping to lead efforts to better understand the role of the environment, diet, drugs and the epigenome on dog aging. 

"I wanted to work at the interface of the public and academic science, and the Dog Aging Project is a great way to do that. So I hopped on the project," said Mccoy, who is a nontraditional student and among the first ASU Online students to pursue a PhD, having spent some time at the National Institute of Aging and Harvard Medical School prior to joining the Snyder-Mackler lab. Mccoy is particularly interested in diet and longevity, and is studying a subset of participating dogs that will be selected to be part of a new clinical study to explore the potential of the drug rapamycin to improve health.

Brassington is a master's degree student now in the molecular and cellular biology program at ASU who hopes to graduate in winter 2022.

"I read about the Dog Aging Project, and I was super excited to find a project that looked at health-related changes due to the environment," she said. She will be examining the environmental air quality and pollution levels of ozone, carbon monoxide and others nationwide. 

Both are now working hard with the citizen science effort, generating experiments, and hope to publish their first data from the Dog Aging Project later this year. 

The Dog Aging Project is actively looking for more participants. If you are a dog owner wanting to get involved, you can learn more at dogagingproject.org.

Written by Jim  Fessenden, UMass Chan Medical School Communications, with contributions from Joe Caspermeyer, ASU.

Top photo: Kristoff, one of Sparky's Service Dogs , keeps watch at the feet of his handler Taylor Randle on Hayden Lawn on Sept. 8, 2016, as they hang out with other puppies being trained by students. Photo by Charlie Leight/ASU News

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The New Science Of Understanding Dog Behavior

Dr. John Bradshaw is an anthrozoologist and a noted scholar of animal-human interactions. Basic Books hide caption

Dr. John Bradshaw is an anthrozoologist and a noted scholar of animal-human interactions.

Dog Sense: How the New Science of Dog Behavior Can Make You a Better Friend to Your Pet By John Bradshaw Hardcover, 352 pages Basic Books List price: $25.99

Read An Excerpt

What's the best advice to give man about respecting man's best friend?

Animal behaviorist John Bradshaw says it's realizing that dogs are neither wolves nor furry humans and that dog owners have certain responsibilities to make sure their dogs are psychologically healthy.

Bradshaw, who has spent much of his career debunking bad advice given to dog owners, is the author of a new behavior guidebook called Dog Sense: How the New Science of Dog Behavior Can Make You a Better Friend to Your Pet . The book details what pet owners should expect from their dogs and what their dogs should expect in return from their owners.

How To Reprimand Your Dog

One of the most common problems owners face, says Bradshaw, is knowing what to do when a dog misbehaves. For example, many owners might be inclined to immediately physically reprimand a dog for jumping up on visitors. But Bradshaw says that's the wrong way to teach your pet how to behave because dogs see any form of attention — even negative attention — as a reward. Instead, he says, owners should immediately ignore their pet completely.

"Most dogs require their owners' attention [and] they want their owners' attention," he says. "They want people's attention in general. And withdrawing that is a very powerful signal to the dog."

Bradshaw recommends folding your arms, looking away and pretending your dog isn't in the same room. Your change in body language will be apparent to your pet.

"Then you'll find that quite quickly the dog begins to realize that [their bad behavior] is not working," he says. "You can then use a distraction technique to get the dog to do something else, like sit or lie down and then it will get the idea that this is what it's supposed to do when visitors come to visit."

Bradshaw says dogs naturally want to please and play with people, especially the people who love them.

"[When a puppy's eyes open it has] a very strong ability to learn about people and ... this behavior persists throughout life," he says. "And surprisingly, most dogs, given the choice, will actually prefer human company to other dog company."

Studies indicate that dogs will naturally gravitate toward humans, though Bradshaw says how that idea gets into a dog's developing brain is still somewhat of a mystery.

"But they have an exaggerated tendency to learn from anything that people do right from the minute they're capable of doing it," he says. "They're particularly sensitive to human body language — the direction we look in, what our whole body language is telling them, pointing gestures. They are much more sensitive to things like that than almost any other species on the planet."

Creating Expectations For Dogs And Owners

Bradshaw says humans also expect dogs to be companionable when they're needed and unobtrusive when they're not. City dogs, he says, are expected to be better-behaved than the average human child and as self-reliant as adults. But these expectations, he says, create problems for modern dogs.

"Many dogs — maybe as many as half the dogs in the West — that are kept in homes have a real problem with being left alone at some point in their lives," he says. "And the problem may last for weeks or years. ... They crave the company of people. They also have a mind which does not have a particularly good sense of time, so when they get left alone, they can immediately begin to think, 'When's anyone coming back? Have I been abandoned forever?' "

Dogs can get extremely anxious as a result, Bradshaw says. But there are bits of training owners can do to help their dogs avoid separation disorders.

"You train your dog to toilet outside. You train your dog to sit on command," he says. "You should also train your dog to cope with being left alone."

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Bradshaw suggests creating a gradual routine when you leave the house: For example, pick up your keys and coat from the same place, in the same way each time you leave. Those behaviors will become triggers for your dog, who will associate them with your impending absence. He also suggests training new dogs by having them become familiar with these actions — and gradually increasing your time away so that your dog has time to become acclimated.

"You go to the door. You come back from the door. You put the coat back on the rack. You put the car keys back down on the shelf. Then you do it again, but this time maybe you open the door," he says. "Then the next time, you go outside the door and come straight back in again. And the next time, go outside and stand outside for 10 seconds. And then come back in again. And what the dog learns at that stage ... the association with you going out and you coming home. And that is enough for most dogs to reassure them — and very quickly you find you can leave them for hours. They've learned that association, and you coming back and making a fuss of them, and so the idea of you going out actually becomes pleasurable instead of something to panic."

John Bradshaw is the director of the Anthrozoology Institute at the University of Bristol in the U.K. He has studied the behavior of domestic cats and dogs for more than 25 years.

Interview Highlights

On common misconceptions about wolves

"The main [myth] ... is that wolves are essentially an intrinsically aggressive animal that is continuously trying to take over whatever group they find themselves in and dominate it. And the new wolf biology really exposed that as an artifact — that particular view of wolves came from wolves in zoos and in wildlife parks, where a bunch of unrelated wolves were basically put together and told to get on with it and, not surprisingly, they got on with it by being aggressive toward one another. The new picture of wolf society is that wolves are harmonious animals. They live in family groups. They get along really well together, and they're almost never aggressive to one another. The aggression comes out when two families meet, so they have very strong family ties."

On playing tug of war with your dog

"Let's take a very simple piece of advice that trainers take out, which is you should never allow a dog to go in front of you through a doorway because it will give the signal to the dog that you are submissive and are therefore allowing him or her — the dog — to become dominant. Take another one. Many trainers advise against playing tug of war games because there is a risk the dog will win and the dog, by winning, will think that you are being submissive and he will therefore be able to control you in the future. We've done research into a number of these things — including the tug of war game — and have shown that the premise is just completely not true. If you do let a dog win over and over again at tug of war, it likes you. It wants to play with you more than it did to begin with because it's having fun. If, on the other hand, the dog gets less attracted to you and doesn't so much want to play with you — again, but there's absolutely no change of the dog's behavior outside of that particular situation of play — the dog does not get into its head that you're kind of a soft touch and that in the future it will be able to control you and whatever you do."

On breeding

"There's still a great genetic variability if you take the dog as a whole. But within a breed, the variation has diminished. So you get all kind of inherited diseases coming up [which are] very difficult to eradicate at the moment while the breed barriers are being maintained."

On military dogs

"I've been involved with training dogs for the military for about a decade now, so I think everybody but me has been surprised by the dog that went in to find Osama bin Laden. They're very valuable dogs. And I must say, if I was in an environment like that, I would actually much rather have a dog ahead of me than another human being because it's another set of senses — and particularly the olfactory sense. These dogs are trained to find and then indicate all manners of things. In that particular instance, it would presumably be explosives and ammunitions and guns and so on."

On dog senses

"They're colorblind to a certain extent but colorblind humans are not that badly handicapped. Their hearing is a little bit more sensitive than ours in the high-pitched region. But it's their sense of smell that really distinguishes them from us. And I don't think we really take up too much recognizance of that. I think dogs have a right to sniff things whenever it doesn't cause a problem to us. When I meet a dog, I hold my hand out. I don't stick my fingers right out, just in case, but I just make a loose fist and put my hand out to the dog. If it's a small dog, I'll squat down. And that dog will want to come and sniff my hand and lick it if necessary. That's a greeting, and I think if we don't do that, I think it's as upsetting to the dog as if we were talking to somebody that we never met before and covered our faces at that point in time, as if we were trying to disguise who we were."

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Perspective article, the new era of canine science: reshaping our relationships with dogs.

• 1 School of Anthropology, University of Arizona, Tucson, AZ, United States
• 2 College of Veterinary Medicine, University of Arizona, Tucson, AZ, United States
• 3 Cognitive Science, University of Arizona, Tucson, AZ, United States
• 4 California State Polytechnic University, Pomona, CA, United States
• 5 Department of Psychology, Western Carolina University, Cullowhee, NC, United States
• 6 Center for Urban Resilience, Loyola Marymount University, Los Angeles, CA, United States
• 7 Animal Welfare Science Centre, Faculty of Veterinary and Agricultural Sciences, The University of Melbourne, Melbourne, VIC, Australia

Canine science is rapidly maturing into an interdisciplinary and highly impactful field with great potential for both basic and translational research. The articles in this Frontiers Research Topic, Our Canine Connection: The History, Benefits and Future of Human-Dog Interactions , arise from two meetings sponsored by the Wallis Annenberg PetSpace Leadership Institute, which convened experts from diverse areas of canine science to assess the state of the field and challenges and opportunities for its future. In this final Perspective paper, we identify a set of overarching themes that will be critical for a productive and sustainable future in canine science. We explore the roles of dog welfare, science communication, and research funding, with an emphasis on developing approaches that benefit people and dogs, alike.

Dogs have played important roles in the lives of humans for millennia ( 1 , 2 ). However, throughout much of scientific history they have been dismissed as an artificial species with little to contribute to our understanding of the natural world, or our place within it. During the last two decades, this sentiment has changed dramatically; canine science is rapidly maturing into an established, impactful, and highly interdisciplinary field ( Figure 1 ). Canine scientists, who previously occupied relatively marginalized roles in academic research, are increasingly being hired at major research universities, and centers devoted to the study of dogs and their interactions with humans are proliferating around the world. The factors underlying dogs' newfound popularity in science are diverse and include (1) increased interest in understanding dog origins, behavior, and cognition; (2) diversification in our approaches to research with non-human animals; (3) recognition of dogs' value as a unique biological model with relevance for humans; and (4) growth in research on the nature and consequences of dog-human interactions, in their myriad forms, from working dog performance to displaced canines living in shelters.

Figure 1 . Canine science is an interdisciplinary field with connections to other traditional and emerging areas of research. The specific fields shown overlap in ways not depicted here and are not an exhaustive list of disciplines contributing to canine science. Rather, they are included as examples of the diversity of scholarship in canine science.

This Perspective represents the final article in a collection of manuscripts arising from two workshops sponsored by the Wallis Annenberg PetSpace Leadership Institute. Leadership Fellows from around the world gathered in 2017 and 2020 to discuss the state of research and future directions in canine science. The individual articles in this collection provide a detailed treatment of key topics discussed at these events. In this final article, we identify a set of overarching challenges that emerge from this work and identify priorities and opportunities for the future of canine science.

The rise of canine science has benefited substantially from public interest and participation in the research process. Unlike many research studies, which unfold quietly in the ivory towers of research universities, the new era of canine science is intentionally public facing. The dogs being studied are not laboratory animals, bred and housed for research purposes, but rather are companions living in private homes, or assisting humans in capacities ranging from assistance for people with disabilities, to medical and explosives detection. Campus-based research laboratories have opened their doors to members of the public who bring their dogs to participate in problem-solving tasks, social interactions, and sometimes even non-invasive neuroimaging studies. Increasingly, dog owners themselves play a significant role in the scientific process, serving as community scientists who contribute to the systematic gathering of data from the convenience of their homes.

This new research model in conjunction with emerging technologies, makes canine science a highly visible field that engages public stakeholders in unprecedented ways. From a scientific perspective, society has become the new laboratory, and in doing so, has facilitated research with dogs of a scope and scale that was heretofore unthinkable. As tens of thousands of dogs contribute to research on topics ranging from cognition and genetics ( 3 , 4 ) to aging and human loneliness ( 5 ), canine science is entering the realm of “big data” and eclipsing many traditional research approaches. Importantly, these advances are occurring simultaneously across diverse fields of science, creating powerful new opportunities for consilience that will make canine science even more valuable in the years ahead. However, maturing this model toward a sustainable future that serves its diverse stakeholders—who include scientists, research funders, members of the public, and dogs themselves—will require careful navigation of key challenges related to dog welfare, science communication, and financial support ( Figure 2 ).

Figure 2 . Visual summary of the key issues identified in this Perspective . A sustainable future in canine science will require (1) research approaches that prioritize and monitor the welfare of dogs, (2) improved science communication to avoid incorrect reporting of study results, and to translate research findings to meaningful change in practices relating to dogs, and (3) availability of research funding that is not tied exclusively to studying the possible benefits of dogs for humans.

Dog Welfare

Globally, animal welfare has been linked to the public acceptability that underpins sustainable animal interactions and partnerships ( 6 ). Where human-animal interactions have failed to meet community expectations, practices and in some case entire industries, have been disrupted or ceased. Recent examples include whaling for profit and greyhound racing ( 6 , 7 ). Science is not exempt from this necessity to meet with public expectations and the new era of canine science must place canine welfare at the forefront. Considering dogs as individuals and co-workers, rather than tools for work or subjects, reflects a community moral and ethical paradigm shift that is currently underway. Reimagining our relationship with domestic dogs in research will also help inform our treatment of other animals. In this way, studies of dogs and our interactions with them can serve as a pioneering new model for many areas of science.

As scientists advocate for the revision of community and industry practices with dogs in light of new evidence, we must apply the same criteria to the conduct of our research. This includes adjusting canine research and training methods to acknowledge the sentience of dogs, and the importance of the affective experience for dogs in both research and community settings ( 8 – 11 ). The discipline of animal welfare science has progressed rapidly over the last two decades, and we have many animal-based, welfare-outcome measures available to us ( 6 , 11 ). Ensuring the well-being of the dogs we study will be as critical to ongoing social license to operate (i.e., community approval) for canine science as it is for working dog interests ( 12 ). Being transparent about the issues of animal consent and vulnerability, as well as offering animals agency with regard to their participation in science are valuable suggestions offered within this special issue. We encourage our colleagues to not just consider this paradigm shift, but to effect it through prioritizing and representing the dog's perspective and welfare in their research.

Although increasingly, researchers may include a single or limited set of canine stress measures in studies exploring dogs' potential benefits to humans, this approach alone does not fill the need for studies that prioritize an understanding of canine welfare as their central focus. Canine welfare should be considered not just as an emergent population-level measure ( 13 ) but rather with respect to the way in which it is experienced: from the perspectives of individual dogs. Commonly used statistical methods from human research, such as group-based trajectory analysis ( 14 ) may offer proven techniques that allow meaningful reporting on populations while reflecting the nuance of shared, sub-group patterns. Such approaches will better reflect individual differences, for example variations in canine personality, social support and relationship styles, as well as other significant factors. One impediment to robust measurement of animal welfare in canine science has been limited funding.

We believe that all granting bodies who fund exploration of the possible benefits to people from dogs should also fund and require the canine perspective to be robustly monitored and reported. Impediments to this work arise not from lack of researcher interest or access to dogs, but rather from challenges to securing funding that is independent from a focus on human health outcomes, or other tangible outcomes of work that dogs perform. To be able to optimize canine welfare, there is an urgent need for increased funding specifically to study the welfare of dogs, in all their diversity. The new era of canine science will identify what dogs need to thrive, propelling us toward a mutually sustainable partnership between people and dogs.

Communication

One area that has not received much attention in relation to canine science is the way in which research findings are communicated outside the empirical literature. Fueled by media reports, interest in canine science and the impact of dogs on human health and well-being has grown substantially in the last 10 years. A survey by the Human-Animal Bond Research Institute found that 71% of pet owners were aware of studies demonstrating that pets improve mental and physical health. Some of these claims are justified. For example, many studies have found that interacting with therapy dogs reduces stress and anxiety and increases positive emotional states in a variety of settings including hospitals, schools and nursing homes ( 15 , 16 ). In other cases, high public expectations about the healing power of pets are not matched by the results of empirical studies. For instance, while the Human-Animal Bond Research Institute survey found that 86% of pet owners believe pets relieve depression, the majority of studies on pet-ownership and depression do not support these conclusions ( 17 ).

Because so many people have extensive personal experiences with dogs, investigators face unique challenges in sharing research results with the public. In their hearts, dog owners believe that their canine companions make them feel less depressed, or that dogs feel guilty when they've eliminated indoors or explored the kitchen garbage—even though research might suggest otherwise. In addition, when it comes to animal companions, people much prefer to read a news article in which visits with a therapy dog improved the well-being of a child undergoing chemotherapy than an article about a randomized clinical trial which found no differences between the well-being of children in a therapy dog group and a control group ( 18 ). Nor is there likely to be much press coverage devoted to methodological issues such as small effect sizes and inappropriate attributions of causality to the results of correlational studies.

Canine scientists and scholars of human-animal interactions (anthrozoologists) are fortunate that the public is intrinsically interested in our research. We feel that it is critical for investigators to make efforts to communicate the findings of important studies to the public. We caution however, that researchers should not overstate the implications of their findings in press releases and conversations with journalists, despite frequent pressure to do so. These distortions could have a negative impact on misleading the public and misrepresenting the actual findings, a problem that is particularly acute in canine science where well-intentioned pet owners may eagerly adopt practices based on media coverage of scientific studies. The now-established discipline of science communication offers guidance for how best to engage with community and research stakeholders in meaningful ways.

Traditionally, science communication has relied on the knowledge deficit model of communication ( 19 ). Directionally one-way, the deficit model operates on the assumption that ignorance is the reason for a lack of community support and application of scientific evidence. Examples where practices have not been updated in response to research findings include dog training methodology ( 9 ) and breeding selection for extreme body types, such as brachycephaly in pugs and bulldogs, even though the health and welfare impacts are scientifically well understood ( 20 ). Scientists who share their research results thinking that knowledge disseminated—to “educate” the public—is enough to result in different dog care decisions, industry practices or legislation, will generally find this to be ineffective ( 21 ). This is because the deficit model overlooks the underlying beliefs, existing attitudes and motivations for current practices. We now recognize that the deficit model is not the most effective way to communicate, engage stakeholders and effect change ( 22 , 23 ).

Further exploration of the effect of targeted and intentional science communication, informed by human behavior change research, will improve the translation of canine science to meaningful outcomes for dogs and people alike ( 12 ). This is important, as many studies in canine science have applied aims designed to inform global policies and the creation of best practices ( 24 , 25 ). Applied research from the livestock and farming sector suggests that coordinating human behavior change strategies from social and psychological sciences can influence beliefs and attitudes to motivate changes in the ways people behave toward animals, resulting in improved animal welfare ( 26 – 28 ). In the era of attention economics, where scientists are competing for public attention alongside other diverse media, it is vital that the communication of our work is honest, relevant, and effective, to ensure that our field stays on the radar of key stakeholders, funding bodies and change agents.

A third key challenge in the future of canine science concerns research funding and a careful balancing of the priorities of scientists and funding agencies. In the last decade, canine science has received considerable support from the pet care sector, as well as human health and defense agencies [e.g., ( 29 )]. Fine and Andersen ( 30 ) stress that although funding is still a challenge in human-animal interaction research, there are now more options to be found. In 2008, the Waltham Petcare Science Institute initiated a public-private partnership with the Eunice Kennedy Shriver National Institute of Child Health and Human Development. Over the past decade, this partnership has provided funding for research aimed at measuring the impact of specific Animal-assisted interventions. Since 2014, the Human Animal Bond Research Institute has funded a total of 35 academic research grants investigating the health outcomes of pet ownership and/or human-animal interaction, both for the people and non-human animals involved. Despite clear benefits for enabling research, there remains a limited group of agencies responsible for funding this work. This has potential to constrain the range of topics being studied. In addition, scientists may feel compelled to support the agendas of industry groups, such as those in the pet sector, who often encourage research that will demonstrate the benefits of pets and human-animal interactions.

These constraints were recognized by Wallis Annenberg PetSpace in 2017 when they envisioned their Leadership Institute Program with a mission to promote interdisciplinary scholarship and convene meetings to accelerate research and policy development ( https://www.annenbergpetspace.org/about/leadership ). This model for engagement inspired the organization to offer two invited retreats (2017, 2020) for a total of 33 experts in the field that provided opportunities for open ended and frank discussion about the nature of human-animal interaction research, and the maturing field of canine science. By providing the space and financial support, plus the opportunity to work together and publish, Annenberg PetSpace provided a way to both illuminate current limitations, and to identify priorities for the future, free of constraints from outside interest groups. These intellectual salons have no specific agenda other than to consider the future of the field and what kinds of questions need to be asked based on what we already know. The results of these two retreats include 14 published refereed papers, plus a suite of collaborations that might otherwise not have happened. We hope that these fellowships and retreats continue and inspire others to support similar initiatives so that scholars across multiple disciplines have the opportunity to experience the transformational exchanges that occur during these programs. The new era of canine science will require diverse funding that is not limited to how dogs can benefit humans, from health, safety and economic perspectives. This change will enable researchers the freedom to further our understanding of dogs and their needs for optimized welfare. In turn, this will allow us to identify how dogs and people can thrive together.

Looking Ahead

We hope that the publications emerging from these retreats will reach a diverse community of stakeholders, including students, early career researchers, animal welfare and advocacy groups, legislators and policy makers, philanthropies, and traditional agency funders. The goal of these papers is to spark imagination for projects not yet engaged and to help set the agenda for future research that can enhance our understanding of human-dog interactions and identify paths to ensure a future of symbiotic relationships between these species.

The vision of this collective group of scholars includes the goal of establishing studies with dogs as a sustainable and broad-reaching research focus. Although dogs provide many advantages as a “model species” —including their phenotypic diversity, and shared environments and evolutionary history with humans—a research model centered around dogs has many additional benefits. Dogs provide a rich, interactive and sentient model with deep implications for the way scientists approach animal research, and animal welfare. Dogs also increase the accessibility of research, both literally, due to their ubiquity and opportunities for large-scale public participation in research ( 31 , 32 ), and figuratively, through a body of work with appeal to the broader public.

The field of canine science has much in common with a similar emerging science, that of urban ecology. Humans are historically at the core of the subject material, but non-human elements are often the focus of the study. As such, the work is always culturally embedded, relevant to a variety of stakeholders, and ultimately expected to improve quality of life. The urban ecologists coined a term Use-Inspired Research ( 33 ) from modifying the existing idea of Pasteur's Quadrant which organizes research questions across the axes of fundamental understanding and considerations of use ( 34 ). Both canine research and urban ecology seek fundamental understanding, but also expect to directly apply the knowledge gained to improve outcomes for their subjects and stakeholders.

By including the public in canine science we not only increase the quantity of the data that we can gather, we serve as ambassadors for a new model of responsible animal research. The result increases the value of human-animal interaction research and creates opportunities for the next generation of interdisciplinary scientists. The goal of this collection has been both to highlight specific recent advances in canine science as well as to identify emerging and overarching issues that will shape the future of this field. The multidisciplinary nature of our work with dogs allows scientists to contribute to a robust research agenda, enhancing our understanding of canines and their impact on society. Ultimately, the nexus of our discoveries should have profound effects on reshaping and enriching our relationships with dogs.

Data Availability Statement

The original contributions presented in the study are included in the article/supplementary material, further inquiries can be directed to the corresponding author/s.

Author Contributions

All authors listed have made a substantial, direct and intellectual contribution to the work, and approved it for publication.

We thank Wallis Annenberg PetSpace for supporting the open-access publishing fees associated with this manuscript.

Conflict of Interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Keywords: canine science, dog, animal welfare, human-animal interaction, science communication, funding, sustainability

Citation: MacLean EL, Fine A, Herzog H, Strauss E and Cobb ML (2021) The New Era of Canine Science: Reshaping Our Relationships With Dogs. Front. Vet. Sci. 8:675782. doi: 10.3389/fvets.2021.675782

Received: 03 March 2021; Accepted: 11 June 2021; Published: 15 July 2021.

Reviewed by:

Copyright © 2021 MacLean, Fine, Herzog, Strauss and Cobb. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY) . The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

*Correspondence: Evan L. MacLean, evanmaclean@arizona.edu

This article is part of the Research Topic

Our Canine Connection: The History, Benefits and Future of Human-Dog Interactions

Dog Breed Doesn’t Affect Behavior, According to New Genetic Research

Personality is shaped by a combination of factors, including a pup’s environment

Margaret Osborne

Daily Correspondent

The science is in: No matter its breed, any canine companion can be a good pupper. 

A new genome study published in the journal Science found breed alone is not an accurate way to predict the personality of your four-legged friend. 

Researchers surveyed the owners of 18,385 dogs, asking questions about their pup’s behavior, such as whether they work at tasks until they’re finished, whether they’re friendly with strangers, or whether they circle before pooping. They also sequenced the DNA of 2,155 pure and mixed breed dogs and compared those to the survey results.

They found breed explains only about 9 percent of the variation in an individual dogs' behavior.

“It’s a major advance in how we think about dog behavior,” Elaine Ostrander, an expert in canine genetics at the U.S. National Human Genome Research Institute who was not involved with the study, tells Science ’s David Grimm. “No breed owns any particular trait.”

You may have heard stereotypes about certain breeds—some, like labs, are more lovable, while others, like chihuahuas, are more aggressive. 

“Any good dog trainer will tell you those stereotypes are a disaster,” Marc Bekoff, a dog-behavior expert at the University of Colorado at Boulder, tells the Atlantic ’s Katherine J. Wu. “Breeds don’t have personalities. Individuals do.”

But breed can tell researchers some things. Overall, the scientists found some behavioral traits are more common in certain breeds. For example, Border collies seem to be more ready to respond to human direction than other breeds. 

"We found things like German shorthaired pointers were slightly more likely to point, or golden retrievers were slightly more likely to retrieve, or huskies more likely to howl, than the general dog population," Kathryn Lord, a researcher at the University of Massachusetts Chan Medical School and author on the paper, tells the New York Times ’ James Gorman. 

Researchers found no behavioral trait was present in all dogs in a breed or missing from all of them either, per the Atlantic .

In other words, though some behaviors are more likely to pop up in some breeds, breed alone cannot predict the disposition of a particular dog. Instead, personality is shaped by a combination of factors, including a dog’s environment. 

"Genetics matter, but genetics are a nudge in a given direction. They're not a destiny," Evan MacLean, the director of the Arizona Canine Cognition Center at the University of Arizona, who was not involved in the research, tells NPR ’s Becky Sullivan. "We've known that for a long time in human studies, and this paper really suggests that the same is true for dogs."

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Margaret Osborne | | READ MORE

Margaret Osborne is a freelance journalist based in the southwestern U.S. Her work has appeared in the  Sag Harbor Express  and has aired on  WSHU Public Radio.

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Dog behaviour has little to do with breed, study finds

Research shows high degree of variability between individual animals – with implications for owners

From sociable labradors to aggressive pitbulls, when it comes to canine behaviour there are no end of stereotypes. But research suggests such traits may have less to do with breed than previously thought.

Modern dog breeds began to emerge in the Victorian era and are often physically distinct – for example, great danes are huge and chihuahuas tiny. But it has often been thought breed can predict behaviour, too.

Now researchers say there’s little sign that’s the case.

Dr Elinor Karlsson of the University of Massachusetts Umass Chan medical school, a co-author of the study, said research revealed a huge diversity of behaviours within each breed.

“Even if the average is different, you’ve still got a really good chance of getting a dog that doesn’t match what people say that breed is supposed to be,” she said.

Writing in the journal Science , the US researchers report how they analysed survey responses relating to the physical traits and behaviour of 18,385 pet dogs, almost half of which were purebred, with genetic data analysed for 2,155 of them.

Analysis of the survey results for purebred dogs suggested about 9% of behavioural variation was explained by breed.

“For the most part, we didn’t see strong differences in breeds, but there are some [behaviours] that are connected to breed more than others,” said Karlsson.

While no behaviour was exclusive to one breed, howling was more common among beagles, while pitbulls and retrievers were more “human sociable”, or comfortable with strangers.

There were also some differences based on dogs’ ancestral functions. For example, herding breeds were, among other traits, more biddable.

But there was a high degree of variability between individuals, meaning it is difficult to predict a dog’s behaviour based on its breed.

To explore whether genetics explained the associations, the team analysed the behaviour of mutts that had different levels of ancestry from particular breeds. The results reveal some traits have a stronger genetic component than others.

Labrador retriever ancestry was associated with mutts who had few qualms about getting wet, yet such ancestry appeared to have no link to human sociability.

“We’d expect that if Labrador retrievers are genetically more human social, we should see that mutts with more Labrador retriever ancestry to be more human social,” said Karlsson.

While the team’s other analyses found human sociability is highly heritable, Karlsson noted the mutt results suggest the genetic variants involved do not appear to be more common in particular breeds. Instead, differences between breeds for this trait may be down to environmental influences, or even owners’ perceptions.

But not all behaviours were found to be heritable, including how easily a dog is provoked by a frightening trigger – a finding that suggests that how aggressive a dog is may have little to do with genetics.

The team says the study has implications for owners.

“They should pay much less attention to all the stories about what their dog’s breed ancestry says about their behaviour and personality, and pay attention to the dog sitting in front of them,” said Karlsson.

Daniel Mills, professor of veterinary behavioural medicine at the University of Lincoln, who was not involved in the work, said genetics can give insights into populations but often reveal far less about individuals.

Mills added that it was not surprising genetics played little role in canine aggression, and criticised breed-specific legislation.

“Potentially risky behaviours … are unlikely to be controlled by simple genetic mechanisms since animals have to make judgment calls based on the much wider environment and their developmental history,” he said.

The findings came as another study, published in Scientific Reports , revealed that different breeds have strikingly different life expectancies.

The analysis of 30,563 records of dog deaths in the UK, collected between 2016 and 2020, showed that while jack russell terriers have a life expectancy of 12.72 years at birth, flat-faced breeds tended to have shorter lives, with French bulldogs having a life expectancy at birth of just 4.53 years.

Such brachycephalic dogs are known to be prone to myriad health problems as a result of their extreme physical traits.

Justine Shotton, president of the British Veterinary Association, said the findings of lower life expectancy for flat-faced breeds was concerning: “While the study doesn’t prove a direct link between these breeds’ potential welfare issues and shorter length of life, the findings serve as a fresh reminder for prospective dog owners to choose a breed based on health, not looks.”

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A Review of Domestic Dogs' ( Canis Familiaris ) Human-Like Behaviors: Or Why Behavior Analysts Should Stop Worrying and Love Their Dogs

Dogs likely were the first animals to be domesticated and as such have shared a common environment with humans for over ten thousand years. Only recently, however, has this species' behavior been subject to scientific scrutiny. Most of this work has been inspired by research in human cognitive psychology and suggests that in many ways dogs are more human-like than any other species, including nonhuman primates. Behavior analysts should add their expertise to the study of dog behavior, both to add objective behavioral analyses of experimental data and to effectively integrate this new knowledge into applied work with dogs.

Dogs in Human Society

“That the dog is a loyal, true, and affectionate friend must be gratefully admitted, but when we come to consider the psychical nature of the animal, the limits of our knowledge are almost immediately reached” – Sir John Lubbock. (1889, p. 272)

Our Intertwined Past

Sir John Lubbock's opinion, outdated though its language may be, is not an inappropriate summary of the state of research on dog behavior today. Domestic dogs are never far away from most people's lives, but objective understanding of their behavior is still surprisingly scarce.

A better understanding of the variables controlling dog ( Canis familiaris ) behavior could have practical importance for the growing number of industries that utilize the behavior of domestic dogs—not only in formal training settings, such as police dogs, drug-sniffer dogs, guide dogs, and so forth—but also in the public realm, where the line between the love of man's best friend and the fear of so-called “bad dogs” is a source of great anxiety. In addition, a more complete understanding of the role of social stimuli, which develops as a result of a natural history of operant and classical conditioning within the domestic dogs' home environment, could play a crucial role in maximizing the quality of our interactions with dogs in a variety of settings.

Humans and dogs share a long intertwined history. DNA evidence suggests domestic dogs most likely diverged from wolves in different places at different times beginning as long as 135,000 years ago ( Vila et al., 1997 ). This is when the morphological structure of certain groups of wolves began to change to more closely resemble the modern domestic dog. Anthropologists and archaeologists have argued that this is an overestimate, claiming that the best way to determine the time of domestication is to look for signs of a close association between dogs and humans ( Morey, 2006 ). One way this has been done is by looking for evidence of dog burials. The earliest burial remains of a domestic dog are 14,000 years old and were found in Bonn-Oberkassel, Germany ( Nobis, 1979 ). The dimensions of the well-preserved lower jaw and teeth suggest that this animal was domesticated and could be compared to a small sheep dog, making it the oldest known domesticated animal and a companion of the Cro-Magnon Man in the late Paleolithic age ( Nobis, 1979 ). The time line of dog burials around the globe indicates the spread of dog domestication at different geographic areas ( Morey, 2006 ).

Role of Dogs in Human Society

The exact location and lineage of the first domesticated dog are still under debate, but the impact that humans have had on the domestic dog as a species is undeniable. Dogs play an astonishing range of roles in human society. Many individuals put their faith in rescue dogs when stranded in the wilderness or capsized in cold water. Others rely on guide dogs to get them safely to multiple destinations on a daily basis. Drug dogs, de-mining dogs, police dogs, termite- and even cancer-detecting dogs are trained and utilized as substance detectors even in the face of competition from the latest technology. There are herding dogs, hunting dogs, sled dogs, and various other specializations that are crucial to the livelihoods of many individuals, not to mention the role dogs play in entertainment and the pleasures of individual dog ownership—sufficiently reinforcing to sustain 74.8 million dogs in the United States, at a cost to their owners of over $100 billion ( American Pet Products Manufacturers Association, 2007 ).

However, qualities desired in one specialization may not be appropriate in dogs filling another capacity. For example, the dependency on human guidance and direction sought in companion dogs may inhibit a rescue dog's ability to problem solve and function independently in situations when its handler is out of sight ( Miklósi, Pongracz, Lakatos, Topál, & Csányi, 2005 ). It is important, therefore, to take breed specializations and individual history into account when selecting dogs for specific tasks. The more that is known about dog behavior, the more that can be done to make the training of working dogs as efficient as possible.

A greater understanding of dog behavior also would be beneficial in a society that perceives dog attacks and consequent deaths to be a growing problem. The Humane Society of the United States estimates that 2% of the population is bitten by a dog each year (over six million people) and ten to twenty of these bites are fatal—with the victim usually a child ( Humane Society of the United States, 2007 ). Recently, the Minnesota Department of Health (2007) reported a 40% increase in the number of hospital treated dog bites between 1998 and 2005. According to attorney Kenneth Phillips this increase in medically treated dog bites is representative of an increase in the dog population at large, which rose 36% from 1986 to 1994 ( Phillips, 2007 ). The public response to increased media reporting of dog attacks has been to label certain breeds as “bad dogs.” Malcolm Gladwell (2006) in the New Yorker likened the profiling of “dangerous dog” breeds to the racial profiling that has dominated the search for terrorists since September 11th, 2001. As with most forms of prejudice and profiling, the banning of specific breeds of dogs from municipalities (most commonly at present the pit bull), fails to effectively identify the environmental causes of undesired behavior so that positive behavior can be reinforced and aggressive behavior controlled with more enlightened methods. Breed profiling may lead not only to a misguided fear of well-behaved dogs identified with a “bad” breed, but may also offer a false sense of security around a dog showing warning signs of aggression just because it comes from a breed with a good reputation.

Phylogeny VS. Ontogeny

Despite the omnipresence of dogs in human lives, scientific study of the factors that have allowed dogs to thrive in human environments has until recently been surprisingly meager.

The causes of the characteristic behaviors of dogs can be understood at two levels. First are the phylogenetic influences on behavior that arise as a result of the unique evolutionary past of domestic dogs. Second, and perhaps more importantly (at least in the sense that they are available for modification in real time), are the ontogenetic causes that are the history of contingencies of reinforcement each domestic dog experiences within human society during its lifetime.

The phylogeny of dogs is particularly interesting because, instead of natural selection by the environment, artificial selection by humans is responsible for the hundreds of breeds of domestic dog that exist today. There is also evidence that selection for desirable physical and behavioral traits has led to many changes in social behavior as unexpected byproducts ( Hare & Tomasello, 2005 ). This has led some scientists to attribute the propensity of dogs for human social interaction to convergent evolution, where the two genetically distinct species were shaped by similar selective pressures ( Hare & Tomasello, 2005 ).

There is, of course, no question that genes play a role in the behavior of domestic dogs, but a dog's individual environmental history plays a major role in shaping its behavior over its lifetime. From the time a puppy is brought into a human household it is completely dependent on human caretakers for all of its needs. The majority of reinforcers a dog will have access to throughout its life are controlled, either directly or indirectly, by humans. This is comparable to the situation of young human children, and may explain in part the similarities in sensitivity to human social stimuli shown by dogs and children. However, unlike children, domestic dogs remain dependent on humans for primary reinforcers, such as food, water, access to mates, and even touch, throughout their lifetimes. Consequently, their access to reinforcers is contingent upon appropriate behavioral responses within the human social environment. Furthermore, behavior directly related to subordinance and dependency is often shaped in dogs from a young age. A puppy that sits by its bowl and whines for food will usually have a greater chance of reinforcement than one who seeks out a source of food on its own, such as from a closet or off a table. Similarly, a dog that gets its leash or goes to the door and barks when it has to relieve itself will likely be praised and be given the opportunity to mark its territory, in addition to lessening the pressure in its bladder. A dog that urinates in the house, in contrast, is likely to receive punishment in the form of scolding and in having its owner clean away its territorial scent. In this way, dependence and sensitivity to human contingencies are shaped quickly in domestic dogs in human households. In many cases reinforcement depends on the dog's ability to recognize social stimuli presented by humans, both subtle human gestures that may serve as discriminative stimuli for certain behaviors and overt mands which command a direct and specific response from the dog.

The Study of Dog Behavior in Historical Perspective

The behavior of dogs was very important in the early history of comparative psychology. Darwin wrote extensively about dog behavior, intelligence and emotions, often using his own dogs as examples. He believed that dogs had emotions such as love, fear, shame, and rage, as well as dreams, and the ability to imitate and reason ( Darwin, 1871 ). Darwin also commented on how domestication impacted the behavior of domestic dogs, decreasing their fear of humans, and he even argued for the evolution of distinct barks with various meanings.

Darwin's neighbor in Downe, Sir John Lubbock, was one of the first to carry out experimental tests of the intelligence of dogs. In the first recorded experiment on nonhuman language abilities, Lubbock trained his dog, Van, to bring him a card labeled “food” by reinforcing this response with the presentation of bread and milk upon retrieval. Once Van could readily discriminate between the “food” card and a blank card, Lubbock added more cards containing words such as “out,” “bone,” “water,” and “tea,” and reinforcing their retrieval with the action or item on the card. Although his data were, by his own admission, preliminary at best, Lubbock reported that out of 113 card retrievals, Van selected the “food” card 80 times and the “tea” card 31 times. Since the dog consumed these items with alacrity, Lubbock concluded that the dog had learned to communicate his wants effectively ( Lubbock, 1889 ). Van's successes inspired Lubbock to attempt to use this method to test the dog's color discrimination abilities as well as its ability to count, but no results were published.

The most famous early researcher to use dogs was, of course, Ivan Pavlov. As is widely known, he discovered the form of conditioning now associated with his name using domestic dogs as experimental subjects. Pavlov exploited this phenomenon to explore dogs' sensitivity to scents, touch, temperature, and musical tones (Pavlov, 1906/1966). Less well known is that he speculated on the role of Pavlovian conditioning in the training of domestic dogs: “You lift the dog's paw saying ‘give me your paw’ or even ‘paw,’ and then give the dog something to eat. After repetition of this procedure the dog gives its paw at these words; it does so without any word of command when it has a keen appetite” ( Pavlov, 1936/1966, p. 309 ).

Current Directions in Dog Research

The last twenty years have seen a resurgence of research into the behavior of domestic dogs. Most of the studies reviewed here have drawn their inspiration from work on developmental and cognitive questions in humans and nonhuman primates. This search for common psychological processes in humans and dogs has been motivated by the fact that humans and domestic dogs have shared a common environment and similar selective pressures for tens of thousands of years.

Responsiveness to Human Social Cues

One of the most interesting behavioral characteristics of the modern domestic dog is its predisposition to attend and respond to human social gestures and cues. Skinner (1953) noted that the behavior of other individuals can be an important source of social stimuli. Gestures and cues are social stimuli that likely started out as behaviors that directly impacted the behavior of another individual in a reinforcing or punishing way. Skinner gives the example of a policeman's “stop” signal, which could have originated from the action of a man putting out his hand against another man's chest forcing him to stop. If this were aversive, the second man might learn to stop before he reached the first man's upheld hand in future presentations ( Skinner, 1986 ). Once a gesture is established, an individual's history with the stimulus shapes his or her behavior in its presence. Thus, if the social contingencies are established already for behavioral responses to a particular gesture, contact with the original behavior that evolved into the gesture is not essential. Other examples of common human gestures include pointing, nodding, reaching towards something, or glancing between an object and another individual. Skinner focused on how gestures might come to act as social stimuli in humans, but the basic principles could easily be applied to dogs as well. For example, if a human throws a dog's ball in a game of fetch, the throwing motion or outstretched arm serves as a discriminative stimulus to chase something in the direction of the release. This reaction most likely ties into the reinforcing effects of chasing a ball or catching prey along with the social reinforcers received for retrieving the object. This behavior, of following the direction of an outstretched arm, may generalize to less dynamic forms of the stimulus, and the dog may begin to follow gestures such as pointing or fake tosses to static objects to be retrieved. The use of gestures could undoubtedly be shaped in many species through this process over time, but dogs appear to demonstrate a sensitivity for human gestures that many other nonhuman species lack ( Brauer, Kaminski, Riedel, Call, & Tomasello, 2006 ; Hare, Brown, Williamson, & Tomasello, 2002 ).

Responses to gestures usually are tested in an object-choice paradigm. In this test, a reinforcing object is hidden in one of two or more locations or containers. The subject enters the test area, and a gesture is given to indicate the location of the object. Alternative hiding places are equated for smell and other cues and are usually sham-baited to control for the effects of noise and human scent. The dog is then allowed to approach the containers and indicate a choice by touching or coming within a required distance of one of the locations.

Miklósi, Polgárdi, Topál, and Csányi (1998) carried out the first study investigating the use of human social cues by domestic dogs. Modeled primarily on studies with humans and nonhuman primates, two bowls were used to hide food items in an object-choice paradigm. One of the bowls was baited out of sight of the subject; the location of the food was determined by a coin toss with no bowl being baited more than twice in a row. The dog then was led back into the room and was held 3 meters from the bowls. The experimenter, who stood behind the bowls, made eye contact with the dog and then gave the predetermined gesture. The experimenter then returned to a neutral position and the dog was allowed to indicate its choice by approaching one bowl. A correct choice resulted in food reinforcement and an incorrect choice ended the trial with no reinforcement. Five gestures were used in this study: pointing, bowing (bending the upper torso), nodding, head-turning, and glancing with the eyes only. Each gesture was presented a minimum of 30 times before the next was introduced. All dogs experienced the gestures in the order given here. To progress to the next gesture, 80% accuracy had to be met for the previous condition.

All 6 pet dogs in this study were able to use pointing, head nodding, bowing, and head turning to identify the target bowl without explicit training in the first fifteen trials, and the only significant initial difference among dogs was the ability to use gazing as a discriminative stimulus ( Miklósi et al., 1998 ). Subsequent studies have analyzed a wider set of gestures and found dogs that can use, or be trained to use, various types of pointing with the arm or extensions of the arm ( Hare & Tomasello, 1999 ; Miklósi et al., 2005 ; Miklósi et al. 1998 ; Soproni, Miklósi, Topál, & Csányi, 2001 ; Soproni, Miklósi, Topál, & Csányi, 2002 ; Udell, Giglio & Wynne, 2008 ), glancing ( Miklósi et al. 1998 ; Soproni et al. 2001 ; Udell et al., 2007 ), local enhancement by a human's presence near the target ( Hare & Tomasello, 1999 ), and a human placing a token on a target ( Hare & Tomasello, 2005 ; Udell et al., 2008 ).

Other studies of the exploitation of human gestures by dogs have varied the way in which conditions are presented. Instead of a hierarchical series of gestures in which one gesture must be learned to criterion before the next is introduced, some studies have presented conditions in orders that vary across subjects and thus control for the effects of generalization from one gesture to another ( Brauer et al., 2006 ). Others have used probe methods to insert novel gestures into a series of trained or familiar gestures such as pointing ( Soproni et al., 2002 ). These studies have largely supported the conclusions from the pioneering work of Miklósi et al. (1998) , suggesting that Miklósi et al.'s positive conclusions were not artifacts of having the dogs master one gesture type before proceeding to the next.

Particularly noteworthy in these studies of dog responsiveness to human gestural cues, is that some of the successful dogs in these studies had only had minimal contact with humans or did not live as pets in human households. Miklósi et al.'s (1998) study included 5 assistant or guide dogs that did not reside in a typical household setting and yet still performed above baseline in the pointing and bowing conditions in the first 15 trials.

The degree to which individual dogs attend to human social cues and their tendency to rapidly integrate new behaviors into their repertoire based on the consequences that follow from them, says something about both their development and their environment. For dogs to provide adaptive responses to human gestures requires not only attentiveness and close proximity to human action, indicative of some sort of social attachment to humans, but also sensitivity to context within a human environment.

This conclusion gains strength from recent research on the role of context in the training of basic commands, such as “sit” and “come.” Fukuzawa, Mills, and Cooper (2005) demonstrated that unintentional human cues influence how dogs respond after training. Dogs who responded to the commands “sit” and “come” reliably when a human was giving the command, showed declines in performance when the command was given by tape recorder in the human's presence, and declined further when the human wore tinted sunglasses. Furthermore, when the human experimenter gave the two commands from behind a screen, out of sight of the dog, the dogs responded dependably to the “come” command but not to the “sit” command ( Fukuzawa et al., 2005 ). This finding makes sense when the context of training and previous exposure is considered. “Come” is often applied and reinforced when presented from a greater distance or when the dog is out of sight, whereas the “sit” command is usually given and reinforced only when the dog is in close proximity to the human issuing the command.

Ability to Cue Humans

Most research in the area of social cues has focused on the dog's response to human gestures, but a study by Miklósi, Polgárdi, Topál, and Csányi (2000) found that dogs that had seen a food item or a toy hidden in a specific bowl placed out of their reach while their owner was out of the room, were able to communicate to their owner the location of the hidden target item when he or she returned. These dogs showed a significant increase in mouth licking, vocalization, sniffing, looking at the owner, and looking at the location of the hidden object after the toy had been hidden and the owner returned. Vocalizations and gaze directed at the location of the hidden object were also higher when the owner was present than when the dog was left alone after hiding, although both behaviors occurred in both conditions ( Miklósi et al., 2000 ). Gazing between the owner and the location of the food or toy occurred an average of three times in the first minute, with 8 out of 10 dogs looking first at their owner and then at the location of the hidden item ( Miklósi et al., 2000 ). This suggests that the dogs remembered where the desired object was hidden after the person who hid it had left the room, and that the dogs displayed behaviors, such as glancing between the naive owner and the location of the object, specifically instrumental in getting the owner to uncover the target object.

Dogs also follow the behavioral cues of other dogs in object-choice tasks. In one study, dogs could find a hidden item at above-chance levels when a trained demonstrator dog oriented towards the correct location while gazing at it, or presented a local enhance cue such as sitting by the correct location ( Hare & Tomasello, 1999 ). The specifics of this study will be discussed later, but these findings may suggest that this kind of “showing” or cuing behavior is part of the everyday behavioral repertoire of the domestic dog.

Object Permanence

“Object permanence” is a Piagetian term for an individual's continued interest in a stimulus after it has disappeared from sight. Children go through several stages of object permanence during development, from a complete disregard for obscured objects at the earliest ages of testing, to sustained search for hidden objects starting around age 2 years. Dogs have been tested for object permanence of varying levels. Gagnon and Doré (1992) completed a series of eight tests with 30 dogs of different breeds. The first four tests made up the visible displacement task. In these tests a toy was placed behind a screen in full view of the dog, and subsequently moved from screen to screen—but always so that the dog could easily observe the movements. The last four tests made up the invisible displacement task. In these tests the toy was first placed in a container before it was moved. The container with toy was then placed behind a screen, the toy was inconspicuously removed from the container and left behind the screen, and finally the now-empty container was shifted behind a different screen. The dog could not know that the container was empty and would naturally go to the container first. An individual that has mastered invisible displacements is one that, on finding the container to be empty, returns to the last place the container stopped to search for the toy. Reinforcement in this study consisted of the opportunity to play with the toy once it was found.

Gagnon and Doré (1992) found that their dogs were more successful on the visible displacement tasks than on the invisible tasks, but some dogs—those that experienced all four visible displacement tasks first before moving on to the invisible displacements—were then successful on the invisible displacement tasks. Unlike the studies of dogs following human gestures, the data in most cases showed improvements across trials rather than a spontaneous ability to follow the cues offered.

This study is important within a Piagetian framework because it showed that domestic dogs could display behaviors characteristic of the sixth, and most advanced, stage of object permanence ( Gagnon & Doré, 1992 ).This conclusion, however, is not universally accepted. Other research has suggested that although dogs still search for the toy in invisible displacement tasks, they are not truly demonstrating stage six behavior because their search patterns do not match those of children in that stage ( Watson et al., 2001 ). Furthermore, a follow-up study provides evidence that dogs in these types of experiments are not showing object permanence at all. Collier-Baker, Davis, and Suddendorf (2004) demonstrated that it was the final resting place of the pole used to move the target ball that cued successful responding, not object permanence after all.

Theory of Mind

Dogs appear to be sensitive to the attentional state of humans and this in turn has an impact on their behavior in a variety of situations. For example, in conditions where taking a piece of food has been forbidden, domestic dogs are much more likely to take the food if the human experimenter does not have a direct view of the food or of the dog approaching the food ( Brauer, Call, & Tomasello, 2004 ; Call, Brauer, Kaminski, & Tomasello, 2003 ). In conditions where the experimenter has a clear view of the dog and the food, dogs typically obey the wait command given by the experimenter. However, dogs instructed not to take the food often disobey if the human's eyes are closed, if the human's back is turned, if the human is distracted, if the human leaves the room, or if some barrier blocks the human's view of the food and the dog's approach to the food ( Brauer et al., 2004 ; Call et al., 2003 ).

Gacsi, Miklósi, Varga, Topál, and Csányi (2004) concluded that the body orientation and eye visibility of a human also has an effect on the begging behavior of dogs. Dogs were given the opportunity to approach and beg from one of two women holding sandwiches. In one condition the two women faced the dog: One with a blindfold on her head, and the other with a blindfold over her eyes. In the other condition one woman faced the dog and tried to maintain eye contact without moving her head or body, whereas the other woman faced away from the dog and ignored it. Dogs were given a piece of food no matter whom they begged from in each trial. Nevertheless, in both conditions, dogs begged significantly more from the seeing or attentive individuals than from the other woman.

Research of this kind often has been used in support of the possibility that dogs possess a “theory of mind” or ability to adopt the perspective of others (e.g., Brauer et al., 2004 ; Gacsi et al., 2004 ). However, such performances also could be due to past experience with similar contingencies. When food is forbidden, taking the food while a human's face is oriented to the food and visible to the dog would likely be punished. However, taking food in situations where the human's face is not appropriately oriented is more likely reinforced by obtaining the food and less likely to be punished. Furthermore, if begging from a person who is looking at a dog usually leads to reinforcement and begging from someone who is not oriented towards the dog does not (as might occur at a family dinner table), then reinforced begging behavior directed towards attentive individuals should increase whereas nonreinforced behavior towards nonattentive individuals should decrease. Despite the fact that reinforcement was available for begging from either woman in Gacsi's study, a long history of begging would increase the probability of begging from an attentive person in the first place, and the dog may never come into contact with the new contingencies. Furthermore, it is possible that a dog reared with different contingencies in place, for example one that only successfully obtains food when humans are not looking, would show the exact opposite behavior. In this case it is not the individual's “theory of mind” which is at stake, but rather the development of a foraging strategy based on the greatest chance of reinforcement.

Word Learning

In 2004 Science published a report of an exceptional border collie named Rico. Rico could recognize vocal labels for over 200 items, mostly toys, which he retrieved by name. Kaminski, Call, and Fischer (2004) demonstrated that Rico also was able to identify a novel item from a group of familiar items and would retrieve the novel item in response to an unfamiliar item name in 70% of trials. Exclusion learning and fast mapping were said to occur, and Rico was credited with not only pairing an unfamiliar name with a novel item but remembering the new name for the novel item in later testing sessions.

Rico's vocabulary is certainly impressive and suggests the potential dogs may have for learning associations between items and human vocal cues. However, Rico's ability to fast map—or pair items with a novel name after one association—was far from perfect. When he was given the name of one of the previously novel items ten minutes after the first pairing, Rico retrieved the correct item out of a pile of nine toys in four out of six trials. However, when tested an hour after the first pairings, using a different set of items, his performance dropped to retrieving the correct item in three out of six trials ( Kaminski et al., 2004 ).

The study with Rico raises interesting questions about the capacity dogs have for name learning and the amount of exposure required to learn the names of new objects. But more research is needed before the implications of his performance become clear. Questions that remain open include: Is Rico an exceptional dog, or could his achievements be replicated in other dogs given appropriate training? What might the necessary and sufficient training regimen be to train a dog to respond to vocal labels in this way? What is the limit of a dog's vocabulary? How does a dog generalize to words pronounced in different ways and by different individuals?

Individual and Breed Differences

It is a widely accepted part of the common lore that different breeds of dogs show characteristic behavioral patterns and aptitudes, and that individual dogs differ in their temperaments. In an attempt to quantify these lay impressions, Svartberg (2004) developed the dog mentality assessment (DMA). The assessment consists of a battery of tests, such as the dog's reaction to social contact, play, chase games, passive situations, strangers, sudden appearances of objects, and loud noises. Factor analysis on these scores led to the identification of six traits: playfulness, chase-proneness, curiosity/fearlessness, sociability, aggressiveness, and distance-playfulness. Follow-up questionnaires indicated the stability and value of the DMA for predicting broader personality dimensions, such as a placement along a shyness–boldness continuum ( Svartberg, Tapper, Temrin, Radesater, & Thorman, 2005 ). However, the DMA has been less successful at identifying and predicting long-lasting aggressiveness and nonsocial behavioral problems ( Svartberg, 2004 ; Svartberg et al., 2005 ).

Evidence for inherent breed and sex differences, as measured with the DMA, is limited. One study indicated that high scores on the boldness scale do not correlate with the breed or sex of a dog. However, breed and sex appear to play some role for the lower scoring dogs, of which female German Shepherds and the other tested breed of working dog, Belgian Tervurens, scored lower than male German Shepherds ( Svartberg, 2002 ). Furthermore, a later study ( Svartberg et al., 2005 ) indicated that scores on all six personality traits did not vary significantly by breed types divided into herding dogs, guarding dogs, and gun dogs. This is not to say that differences do not exist between individual breeds; however, the source of these differences does not seem to be explained by the current personality tests or models. More tightly controlled behavioral methods, focusing on individual differences and specific behavior problems, may have more success at getting to the root of the environmental influences that shape these differences in behavior.

Social VS. Causal Cues: The Difference Between Dogs and Nonhuman Primates

Ever since Darwin (1859) , the search for human-like social cognition (i.e., behavior controlled by human and conspecific social cues similar to that observed in humans) has focused on our closest genetic relatives, particularly chimpanzees. Though much remains controversial in this field, it seems clear that chimps and several other species of primates are only modestly successful on many tasks designed to test for human-like social reasoning. Thus, chimpanzees are only able to follow gaze and show joint attention under a limited set of conditions ( Barth, Reaux, & Povinelli, 2005 ). In the object-choice task described above, few chimpanzees or other nonhuman primates are able to use gaze or other social cues such as pointing to identify the location of a hidden object ( Call, Hare, & Tomasello, 1998 ; Call & Tomasello, 1998 ; Itakura, Agnetta, Hare, & Tomasello, 1999 ; Povinelli, Reaux, Bierschwale, Allain, & Simon, 1997 ; Tomasello, Call, & Gluckman, 1997 ). Successful individuals typically need dozens of repeated exposures to the cue, and show poor transfer after even small changes to the testing environment ( Brauer et al., 2006 ; Call, Agnetta, & Tomasello, 2000 ; Itakura et al., 1999 ).

Dogs, in contrast, though they share much less of our genetic material than do chimpanzees, nonetheless show a spontaneous ability to follow human gestures to find reinforcing objects, even in the absence of training in the laboratory. Most remarkably, even dogs raised with minimal human contact can follow a human point and gaze gesture without explicit training ( Hare et al., 2005 ).

Chimpanzees also have been the species most intensely studied for any ability to respond to the attentional state of humans or conspecifics—so-called “Theory of Mind” abilities. However, several published studies have failed to find any evidence of a sensitivity to another's knowledge (e.g., Brauer et al., 2006 ; Povinelli & Eddy, 1996 ), and studies that do suggest this ability (e.g., Hare & Tomasello, 2004 ) have been subject to extensive criticism (e.g., Boesch, 2007 ; Heyes, 1998 ; Penn, Holyoak, & Povinelli, in press ). Dogs, in contrast, respond readily to human cues in these kinds of tests (e.g., Brauer et al., 2004 ; Call et al., 2003 ; Gacsi et al., 2004 ).

Chimpanzees have been by far the most intensively studied species for the comprehension of human language (including seminal studies by Gardner & Gardner, 1969 ; Savage-Rumbaugh et al., 1993 ; Terrace, 1979 ), but no peer reviewed paper has ever claimed the rapid “fast-mapping” of language acquisition found by Kaminski et al. (2004) in the dog Rico.

Several theories have been proposed to explain why dogs perform so well on tasks involving socially mediated stimuli. The possibility that dogs learn to attend to human social cues simply because of the intensity of their interactions with humans appears to be refuted by the observation that even puppies and domesticated fox kits that have had only minimal exposure to human beings, nonetheless respond very accurately to human cues in choice paradigms ( Hare et al., 2005 ).

Hare and Tomasello (2005) considered the possibility that domestic dogs' high sensitivity to social cues is an evolutionary legacy inherited from wolves, the dog's closest wild relative and progenitor. If general social traits common to wild canids have simply been inherited by domestic dogs, then wolves also should do well on tasks involving social cues. However, when compared to wolves and wild foxes, domestic dogs (including puppies) make significantly more correct responses on choice paradigms where social cues serve as the discriminative stimuli ( Hare et al., 2002 ; Hare & Tomasello, 2005 ). This is true even though the wolves tested had been socialized and raised by humans in their homes as pets. Thus, it does not seem that domestic dogs simply inherited the predisposition to attend to social stimuli from wolves.

Hare and Tomasello's (2005) study included, alongside tests on domestic dogs, comparison tests on fox kits that had been selectively bred over 46 years for nonaggressive behavior towards humans. These fox kits were compared to others reared under the same conditions but not selectively bred for low aggression. Neither group of foxes had been raised in human homes (nor had the earlier generations from which they were descended). Hare and Tomasello found that the fox kits bred for nonaggressive reactions to people performed just like domestic dog puppies on pointing and gazing tasks. The fox kits that had not been selectively bred performed poorly on these tasks, at a level similar to that seen in wolves ( Miklósi et al., 2003 ).

These results suggest that during domestication, traits that were often selected by humans, such as lack of aggression and fearlessness towards people, may have carried with them other genetic traits that led to a heightened responsiveness to human social stimuli ( Hare et al., 2002 ; Hare & Tomasello, 2005 ). It also is possible that by removing genetic tendencies towards aggression and fear towards humans, other preexisting social behaviors were no longer blocked and thus could increase in frequency.

If selective breeding and domestication serve as a likely explanation for the success of domestic dogs on tasks involving human social cues, then that begs the question—Why don't other domesticated animals share these abilities? In fact, domestic cats have been shown to be only slightly less successful than dogs in using basic pointing cues to find a hidden food item in a simple choice test ( Miklósi et al., 2005 ). However, when presented with an unsolvable task, where food was hidden in a butter pot but tied to a stool in such a way that retrieval was impossible, dogs looked between the problem and their owner more often and for longer periods of time, whereas cats only occasionally looked towards their owners and spent much more time trying to get the food themselves. This may indicate that: (1) During domestication cats were selected for traits less tied to the approach of humans and fear reduction, or (2) less stringent contingencies exist for cats in their home environment leading to behavior that is more independent of human action, or both.

The lower responsiveness and less frequent orientation of cats to human cues may in fact be related to the fact that domestic cats are closer to their wild relatives than dogs are to wolves. The domestic cat ( Felis catus ) shows only a low level of genetic divergence from its two nearest wild relatives (the European wildcat, F. silvestris , and African wildcat, F. libyca ), and the earliest evidence for cat domestication is only around 8,000–9,500 years BP—considerably more recent than that for dogs (between 14,000 and 135,000 years BP) ( Driscoll et al., 2007 ; Serpell, 2000 ; Vigne, Guilaine, Debue, Haye, & Gérard, 2004 ). The traits selected for in the domestication of the two species also may have led to differences in the responsiveness and attentiveness each has towards humans. Even today, many dog breeds are selectively bred to work in close association with humans, filling specific roles in industries such as farming, therapy, police, and search- and-rescue. Even with earlier partnerships such as hunting it is quite probable that a dog that stayed close to its owner or was quick to respond to its owner's actions would have been a more beneficial working companion, securing its place in the group and ultimately in the gene pool.

Cats were likely used as mousers and kept as pets from early in their domestication ( Vigne et al., 2004 ), but they are not typically bred for purposes that require a close partnership with humans, even today. Thus, a house cat's independence could have actually been a beneficial trait that increased the chances of its survival in the same environment. Furthermore, cats are often chosen as pets because they are considered low maintenance compared to dogs. They do not require walking, they sleep or entertain themselves most of the day, and they are typically small and quiet enough to go unnoticed much of the time. Thus, there are many more opportunities for cats to engage in independent behaviors without immediate human consequences within the home environment.

Several studies have looked for key similarities and differences between wolves and dogs. Perhaps the most striking developmental difference between dogs and wolves is that, whereas dogs can be socialized to humans within the first sixteen weeks of life, wolves must be removed from their mother for human socialization before fourteen days of age, or acceptance of humans is very unlikely ( Klinghammer & Goodman, 1985 ).

Implications for Phylogeny and Ontogeny

In a study by Frank and Frank (1982) , domesticated dogs (Alaskan malamutes) and wolf pups that were raised in identical conditions in a home environment showed distinct differences in both physical and social development. Conducted as a two-stage experiment, 2 malamutes acquired at 10 days old were compared to 2 wolf pups acquired at 11 days old a year before. The wolf and dog pups did not interact, but the conditions were kept almost identical for the two groups during the experiment. Interestingly the two major differences were that wolves were given more socialization to humans, as they were required to sleep with their human foster parent two out of every three nights as pups, and the malamutes, who did not receive this extra socialization, were given slightly more frequent exposure to the outdoor enclosure. All pups were nursed by the same wolf mother until weaning, at which point they were hand raised and fed by humans.

The wolves reached several physical developmental landmarks days ahead of the malamutes. For example, the wolf pups began climbing over their 45-cm pen wall at only 19 days, whereas the malamute pups could not climb over their 15-cm den box opening at 32 days old. However, socialization of the wolves was much more difficult than of the malamutes. At 2 weeks of age the wolf pups avoided the human handlers whenever possible and hid behind the wolf dam when humans approached. At 6 weeks they became less fearful but somewhat indifferent to the human presence, preferring to be around adult wolves or dogs in the enclosure. The malamutes, in contrast, became more independent of the nursing wolf, and actively approached nearby humans and engaged in “greeting frenzies” on a regular basis ( Frank & Frank, 1982 ).

However, this study has some potential flaws. First, all of the pups were raised by a wolf foster mother, which could have potentially impacted the behavior of the mother to the foster pups or the behavior of the growing pups toward the foster mother. Without a comparison using a Malamute foster mother for both species it is impossible to say that having a same-species foster mother would not produce a closer bond to that individual and therefore less of a bond towards humans. Second, since the two groups of pups were raised at different times, other factors may have been present in one study that were not accounted for in the next, for example, the age of the foster mother or other canine group members and the previous experience of the experimenters raising wolfs before raising the Malamutes.

To address some of these concerns, Kubinyi, Viranyi and Miklósi (2007) conducted a similar study comparing the development and behavior of wolf pups and mongrel dog pups in foster homes with human caretakers. In this study, all pups were individually assigned to a human caretaker who hand raised and fed his or her pup from 4 to 6 days old. Both sets of pups participated in multiple behavioral tests from 3 to 9 weeks of age. When the wolves reached 9 weeks of age they had to be integrated into a captive wolf pack, but were still visited by their caretakers at least once or twice a week. Unfortunately the mongrel dogs in the study continued to live in a human household at this point, so testing later in their lives could have been impacted by different home environments. Nevertheless, the study found that the wolves could be handled by their caretakers similarly to dogs when tested between 1 and 2 years of age. This included coming when called, sitting and lying down on cue, allowing dog accessories such as a muzzle to be put on, and minimal social and physical neophobia. The level of attachment, measured by the length of time the wolves spent in close proximity to their caretaker at 1 to 2 years of age, however, was less for wolves than it was for the dogs. The domestic dogs also out-performed the wolves on tasks involving more complex human social cues, such as momentary distal pointing. The wolves could be taught to use the same level cues as the dogs at 11 months, but only after extensive training ( Kubinyi et al., 2007 ).

Studies comparing domestic animals and their closest genetic relatives are a good step in the direction of identifying the role phylogeny and ontogeny play in key behaviors that seemingly make the species behaviorally distinct. However, much care needs to be taken to make sure both species are treated equivalently and that the behavior that results is not a byproduct of some unintended aspect of the experimental environment. This includes taking into account genetic and developmental differences that may impact how different species respond to stimuli when presented at the same age or in different environments.

The fact that various domesticated animals do better than their nondomesticated relatives on tasks requiring the use of human social stimuli indicates that selective breeding and domestication play some role in this class of behavior. These genetic traits or predispositions may have been a result of artificial selection in some species, but they are still a product of the evolutionary history of that species. Instead of mountains creating the geographic isolation of a pack of wolves, stone walls and chains may have determined which individuals could breed. In place of a natural distribution of ecological resources, a human hand may have determined which individuals would live or die within a pack.

Dogs may have developed at least some behaviors similar to those of humans because the two species lived in such close proximity over 10,000 years. It also is the case that it would have been beneficial to humans to create similar or complementary social traits in these animals through selective breeding. Of course, over most of this history of artificial selection, the human breeders would have understood nothing of genetics or selective breeding. Simple operant conditioning would be sufficient to explain the selection of dogs with desirable traits. Dogs that bit or attacked a human may have been killed, whereas ones that worked well with humans on the hunt and were nonviolent to their owners were taken care of and had a greater chance of reproductive success. Over time, people would have learned to recognize traits in puppies that had typically led to aggressiveness in older dogs in the past, and the process of selecting desired individuals and rejecting ones with undesirable traits would have become more efficient. In other words, the selection of particular traits in dogs would be reinforced with the presence of cooperative, nonaggressive dogs, whereas the tolerance or selection of other traits might be punished with aggressive attacks or a lost investment of food and energy if a fearful dog runs away.

Of course, phylogeny may set the limits of what is possible in behavior, but it is ontogeny—the personal history of reinforcement—that determines what an animal actually does. In a study by Hare and Tomasello (1999) , domestic pet dogs demonstrated the ability to use the location and gestures of both humans and other dogs to help locate hidden food. Four conditions were used: human–local enhance (the human squatted by the correct location); dog–local enhance (another dog sat by the location); human–gaze- and-point; and dog–gaze-and-point (the other dog faced and looked towards the location). When performance was assessed as a group, the 10 subject dogs in the study found food significantly more often in each of the experimental conditions than in the control or baseline condition where no cue was provided. As a group, no one condition appeared to be more helpful than another. However, individual dogs differed greatly in which stimulus they were most successful in using to find the target location. Only 2 dogs were successful in all four conditions, 1 dog was successful in three conditions, 2 dogs were successful only with the human communicator, 2 only with the dog communicator, 1 during both the human– and dog–local enhancement conditions only, and 2 during the human–local enhancement condition only. These differences are most likely due to different levels of experience in the home with situations similar to the ones the experimenters set up in the laboratory.

Dog Profiling Revisited

If, then, there is a genetic component to some aspects of behavior that have a clear impact on human–dog interaction, can bans targeting “bad dog” breeds such as pit bulls, or profiling based on genes in general, be justified by maintaining the position that behavior is a product of genetic tendencies as well? Evidence suggests that the answer is no. Although bites and deaths attributed to pit bulls are up in recent years ( Sacks, Sinclair, Gilcrist, Golab, & Lockwood, 2000 ), other breeds have been number one for aggression against humans at other times. German shepherds and St. Bernards were estimated to be responsible for the majority of deadly dog attacks, not including police dogs, from 1975 through 1980 ( Pinckney & Kennedy, 1982 ). In the 1970s, Dobermans were on the top of the list (Randall Lockwood of the ASPCA, as cited in Gladwell, 2006 ), and between 1993 and 1998 Rottweilers were the most dangerous dog breed ( Sacks et al., 2000 ). However, these estimates are imperfect because they do not take into account the baseline populations of each breed in the U.S. at any given time, and identifying an individual as a specific breed is not always clear cut. Therefore, breeds that have a larger population may be involved in more attacks than less popular breeds but proportionally may be less aggressive; and aggressive dogs that do not fall clearly into a breed category are often labeled as a breed that is already deemed aggressive, thereby inflating the numbers for that breed. However, even in times where one breed may show proportionally higher levels of aggressive behavior, there is evidence that this is not solely due to an inherited “bad dog” gene. In fact, the type of owner, not the breed of the dog, is the best predictor for dog attacks ( Gladwell, 2006 ; Siebert, 2004 ). In a quarter of fatal dog attacks, the owners previously had been arrested for illegal fighting, and many aggressive dogs are ones that have been abused, starved, or deprived of medical attention. In addition, some owners seek out breeds that have a reputation as “bad dogs” and then shape the aggressive behaviors that later seal their fate. According to Randall Lockwood, a senior vice-president of the ASPCA, “A fatal dog attack is not just a dog bite by a big or aggressive dog. It is usually a perfect storm of bad human–canine interactions—the wrong dog, the wrong background, the wrong history in the hands of the wrong person in the wrong environmental situation” (cited in Gladwell, 2006 , p. 26).

Dogs may become problems in human society because their owners may also respond in unconventional ways to social stimuli within the environment because of their own history—possibly exposing their dog to contingencies governed by an abusive, isolated, or neglectful home environment. Therefore, to fully address these and other types of behaviors demonstrated by domestic dogs, the specific contingencies that surround the operant and the specific properties of social stimuli that serve as effective discriminative stimuli need to be identified and defined.

Dogs: Our Closest Relatives?

It may sound strange, but it is not unreasonable to view dogs and humans as subject to convergent evolution ( Hare & Tomasello, 2005 ). Over the last 100,000 years, the social environments of domestic dog pups and human children have become more and more similar to each other, and less like those of either species' closer genetic kin.

It is as a consequence of this intense cohabitation that dogs have come to emulate some behaviors that are commonly viewed as uniquely human, such as the recognition of another's attentional state. These kinds of complex behaviors are commonly structured in relatively vague cognitive terminology. We hope this review will inspire behavior analysts to use the empirical tools of our field to investigate just how closely dog social behavior maps onto human use of social cues. Such research could answer fascinating questions in the evolution of complex behavior, as well as enabling us to live more safely and profitably with our “best friends.”

To Shape, Lead, Catch, or Click?

The study of dog behavior may seem new to experimental behavior analysis, but the interest in applying behaviorist technology to dog training dates back to Skinner's own writings. Skinner wrote: “Since nearly everyone at some time or other has tried, or wished he knew how, to train a dog, a cat, or some other animal, perhaps the most useful way to explain the learning process is to describe some simple experiments which the reader can perform himself” (Skinner, 1951/ 1999, p. 605 ). He went on to provide techniques to shape the behavior of any animal the reader could “catch” using the basic principles of positive reinforcement (Skinner, 1951/1999). Three decades later, Karen Pryor reintroduced behavioral methods of dog training to a new generation of animal trainers and pet owners ( Pryor, 1984 ).

Notwithstanding Skinner's and Pryor's encouragement to behavior analysts to become involved in dog training, and even a paper in the psychological literature calling on behavioral scientists to become more involved in the scientific development of dog training methods ( Tuber, Miller, Caris, Halter, Linden, & Hennessy, 1999 ), the two flagship journals of the field, the Journal of the Experimental Analysis of Behavior and the Journal of Applied Behavior Analysis , have published surprisingly few empirical papers involving dogs as subjects, the most recent ( Cohen, 1970 ) having a publication date of over 35 years ago.

Surely there would be no better way to convince people of the effectiveness of scientific behavioral techniques than to provide them with the technology that they desire to address their current needs. Now is the time to provide the research that will help behaviorists, and in turn society, better understand the behavior of domestic dogs as a species, and to devise refined and easily applied methods of training and evaluation grounded in empirical and testable approaches to behavior.

Acknowledgments

We thank Ray Coppinger, Leonard Green, Bill Roberts, and John Staddon for most helpful comments on a previous draft.

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New Study Shows Importance of Understanding Dog Behavior

A study of pain shows why one should become "fluent in dog".

Posted March 21, 2018

The importance of knowing your dog as the individual they are

I'm a fan of all people who choose to bring a dog into their homes and hearts taking the time to become amateur ethologists and spending time becoming "fluent in dog ." This really isn't asking too much, because when we make this decision we become their caregivers and they assume we have their best interests in mind from "cradle to grave," the cradle begins when we welcome them into our lives.

Learning about dog behavior, even some of the basic rudiments of why they do what they do, is not only fun, but also can be used to know how they're feeling. It's also an excellent way to learn about individual behavioral variability, even among littermates, and to use this information on the individual's behalf. Those who carefully watch, train, and treat dogs for various psychological and medical conditions know that there is no individual being we can accurately call "the dog," and generalizations often fail when an individual's background and personality are ignored (for more discussion please see " My Own Dog Is an Idiot, but She’s a Lovable Idiot " and Canine Confidential: Why Dogs Do What They Do ).

Because of my own interests in "all things dog" and behavior in particular, I was very pleased to learn of a new study by Ana Luisa Lopes Fagundes and her colleagues called " Noise Sensitivities in Dogs: An Exploration of Signs in Dogs with and without Musculoskeletal Pain Using Qualitative Content Analysis. " The entire piece is available online as is a brief and adequate easy-to-read summary titled " Dogs with noise sensitivity should be routinely assessed for pain by vets. " The latter essay begins, "Dogs which (sic) show fear or anxiety when faced with loud or sudden noises should be routinely assessed for pain by veterinarians, a new study has found." The noises that can trigger behavioral responses such as fear and anxiety range from "fireworks, thunderstorms and aeroplanes, to gunshots, cars and motorbikes."

To study the relationship between noise sensitivity and pain, the researchers examined the clinical records of 20 dogs at the University of Lincoln (UK). The data set was comprised of 10 “clinical cases” of dogs exhibiting neuromuscular pain and 10 “control cases” of dogs who did not show any pain. Both groups were similar in breed and age. It's hypothesized that noises that make dogs startle may cause muscles to tense and this can exacerbate pain. Lead researcher Ana Luisa Lopes Fagundes notes, "The aim of the study was to explore the presenting signs of dogs with generalised noise sensitivity with and without pain in their muscles or joints. We think that dogs with this sort of chronic pain may experience the noise quite differently, because if the noise makes them startle it may cause them to tense their muscles and as consequence they feel pain associated with the noise."

Age of onset of behavioral responses to pain is important to consider. The researchers learned that the average age of onset of noise sensitivity was nearly four years later in the “clinical cases.” They write, "This strong theme of an older age of onset suggests that the pain may develop later in life and that owners seek treatment more readily, perhaps because the appearance of the problem is out of character in the subject." Knowing this means that the humans have a good idea of what is typical behavior for their dog(s), and this means that they have previously watched them carefully.

The researchers also note that one marker of pain is that dogs might generalize noise sensitivity to a wider environment and this might prompt their humans to seek medical attention . In the summary we read, "In both cases, the presenting signs of the dogs' behavioural issue included shaking, trembling and hiding, but those with a diagnosed pain issue also showed a higher level of avoidance when it came to places they had a bad experience with noise - for example attempting to avoid a certain area at a park altogether compared with those without pain." Dogs in pain also avoided other dogs.

Becoming fluent in dog: The importance of knowing your dog and watching them carefully

Clearly, to know what a dog is feeling it is essential to know them as an individual . What's a loud or disturbing noise for one dog might not be for another dog. Among the many dogs with whom I share my home, there was great variation. A couple were truly scared of thunder, whereas some didn't show any response at all. One dog trembled when there were sirens, while others weren't affected at all. I did note that as my canine companion Jethro aged, he became more sensitive to sounds, and I knew that he was suffering from a neuroma near the base of his tail. However, I never thought that his heightened sensitivity to noises might have been related to the pain from which he suffered but didn't display behaviorally. Along these lines the researchers note, "It is also possible that the presence of a musculoskeletal pain focus and sound sensitivity interact to lower reactivity thresholds to related stimuli."

When people take the time to become amateur ethologists and citizen scientists they can acquire skills that can truly benefit the dogs with whom they share their lives. It's a win-win for all, and the current study shows just how important it is to pay careful attention to changes in behavior because they can be reliable indicators of pain that might otherwise go undiagnosed. And the good news, according to the researchers about whose work I'm writing, is, "Prognosis seems to be excellent if the case is properly managed following the identification of the role of pain."

Far too many dogs don't get what they want and need in a human-dominated world (for more discussion please see " Dogs Want and Need Much More Than They Usually Get From Us "). They depend on us to know what they want and what they need, perhaps especially when they're suffering and in pain. We are obliged to do so.

Bekoff, Marc. Canine Confidential: Why Dogs Do What They Do . Chicago: University of Chicago Press. 2018.

Ana Luisa Lopes Fagundes et al, Noise Sensitivities in Dogs: An Exploration of Signs in Dogs with and without Musculoskeletal Pain Using Qualitative Content Analysis , Frontiers in Veterinary Science (2018). DOI: 10.3389/fvets.2018.00017

Marc Bekoff, Ph.D. , is professor emeritus of ecology and evolutionary biology at the University of Colorado, Boulder.

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The scholar’s best friend: research trends in dog cognitive and behavioral studies

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• Published: 21 November 2020
• Volume 24 , pages 541–553, ( 2021 )

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• Massimo Aria 1 ,
• Alessandra Alterisio 2 ,
• Anna Scandurra 2 ,
• Claudia Pinelli 3 &
• Biagio D’Aniello   ORCID: orcid.org/0000-0002-1176-946X 2  

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In recent decades, cognitive and behavioral knowledge in dogs seems to have developed considerably, as deduced from the published peer-reviewed articles. However, to date, the worldwide trend of scientific research on dog cognition and behavior has never been explored using a bibliometric approach, while the evaluation of scientific research has increasingly become important in recent years. In this review, we compared the publication trend of the articles in the last 34 years on dogs’ cognitive and behavioral science with those in the general category “Behavioral Science”. We found that, after 2005, there has been a sharp increase in scientific publications on dogs. Therefore, the year 2005 has been used as “starting point” to perform an in-depth bibliometric analysis of the scientific activity in dog cognitive and behavioral studies. The period between 2006 and 2018 is taken as the study period, and a backward analysis was also carried out. The data analysis was performed using “bibliometrix”, a new R-tool used for comprehensive science mapping analysis. We analyzed all information related to sources, countries, affiliations, co-occurrence network, thematic maps, collaboration network, and world map. The results scientifically support the common perception that dogs are attracting the interest of scholars much more now than before and more than the general trend in cognitive and behavioral studies. Both, the changes in research themes and new research themes, contributed to the increase in the scientific production on the cognitive and behavioral aspects of dogs. Our investigation may benefit the researchers interested in the field of cognitive and behavioral science in dogs, thus favoring future research work and promoting interdisciplinary collaborations.

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Introduction

The domestication of wolves was probably the first human successful attempt aimed to control an animal. After the first stage, in which primitive dogs were domesticated from their wild ancestors—the wolves, dogs were artificially selected resulting into the modern breeds, based on different specialization and morphology (Wayne and Ostrander 2007 ). However, several aspects of this history remain unclear, despite scientific efforts in studying dog evolution. Under dispute is the place of the domestication process of dogs—from Europe or southern East Asia—as uncertainly lies in the divergence between wolves and dogs (Thalmann et al. 2013 ; Wang et al. 2016 ). Many doubts also surround the question of how domestication of dogs began and how this process has impacted cognition and behavior in dogs (Hare et al. 2002 ; Udell and Wynne 2008 ; Wynne et al. 2008 ; Topál et al. 2009a ). Probably, the secret of such a long duration and effective cooperation with humans are dogs’ advanced social skills, which allowed them to exchange communicative signals effectively with the human (Miklósi 2009 ; D’Aniello and Scandurra 2016 ; D’Aniello et al. 2017 ; Scandurra et al. 2017 , 2018 ).

In history, dogs were mostly employed for utility, whereby the term “man’s best friend” originated in the eighteenth century (see Miklósi and Topál 2013 ). They are now increasingly involved in different working and sporting activities, and their presence in our homes as pets is also in a growing trend in many countries (see, for example, Murray et al. 2015 ). Pet dogs could promote the welfare of the human family they live with, whereas working dogs are an integral part of social functioning. Several studies demonstrated that keeping dogs has a positive effect on our physical and mental health (Levine et al. 2013 ; Ownby et al. 2002 ; Raina et al. 1999 ; Kramer et al. 2019 ). Although this so-called “pet effect” (Allen 2003 ) received some criticism, noting that some papers reported null or also negative effects on health and happiness of pet owners (see Herzog 2011 for a review), it contributed to the flurry of research on dogs.

Altogether, dogs have become an important social phenomenon attracting scientific interest (Morell 2009 ). There are different reasons why canine research is advantageous beyond the easy access to the subjects for experimental purposes. They show many similarities with humans (Scandurra et al. 2020 ) and, therefore, can be used as a model for human studies. Indeed, there are functional parallels in a range of behavioral features, which are not shared with the closest human relatives, the great apes (Topál et al. 2009b ). The success of dogs as behavioral models also relies on their origin from ancestors with high social behavior, the adaptiveness in living in the anthropogenic niches and the socialization with humans during ontogeny (Kubinyi et al. 2009 ). Moreover, dogs have also been used as a model for comparative and translational neuroscience, cancer and cognitive decline, such as in Alzheimer’s disease in humans (Head et al. 2000 ).

Bibliometric research focusing specifically on the dog’s personality or temperament showed about 50 papers in the database between 1934 and 2004 (Jones and Gosling 2005 ). A comprehensive study by Bensky et al. ( 2013 ) aimed at identifying the major trends in the literature related to the areas of cognitive research on dogs, detected an increase in the studies over the 15 years before the date of the review. However, a worldwide trend of scientific research on dog cognition and behavior has never been explored to date using a bibliometric approach (Chen 2003 ), while the evaluation of scientific research has increasingly become important in recent years. Bibliometric analysis is a useful tool to measure the output of scientific research, using specific indicators to obtain information about the research trends in different fields (De Battisti and Salini 2013 ; Wallin 2005 ).

There are pure cognitive studies, which analyze the brain functioning through brain imaging (e.g., fMRI studies), without taking into account the behavioral responses. However, most of the papers dealing with cognition also include behavioral outcomes. Indeed, brain imaging studies often analyze brain functioning, while the experimental subjects are performing various behavioral tasks. Moreover, many studies provide data for the understanding how stimuli are processed (i.e., cognition) by studying behavioral responses. Therefore, it is not so common to find pure cognitive or behavioral studies, whereby, in this paper, we have considered all studies dealing with both cognition and/or behavior.

Since there is evidence of a growing trend in scientific production (Fanelli and Larivière 2016 ), the first goal of the present paper was to verify whether the dog cognitive and behavioral studies show a growing trend exceeding those of cognitive and behavioral sciences in general. To this scope, we have provided a comparison between the trend in the literature on dogs and that of the whole collection of studies in the subject category “Behavioral Sciences” from 1985 (i.e., the year of starting electronic access to Web of Science database) to 2018. It was verified that peer-reviewed publications on dog cognitive and behavioral studies showed a steeper growth curve with respect to that of the subject category “Behavioral Sciences” starting from 2005. Therefore, this year was chosen as a “starting point” for a “recent analysis” until 2018, including 13 years. To further emphasize the more current changes in the scientific production related to the cognitive and behavioral studies on dogs, we compared the “recent analysis” with an “earlier analysis”. The latter covered in the backward direction an equivalent number of years to the “recent analysis” from the “starting point”.

Our second goal was to understand whether the growth of scientific production in dog cognition and behavior was simply related to an increased research effort in the same research themes or changes in research themes and the contribution of new research themes to this trend.

The further aim was to provide a bibliometric analysis related to sources, countries, affiliations, co-occurrence network, thematic maps, collaboration network, and world maps of the scientific activity related to the cognitive and behavioral studies on dogs. We also attempted to identify the most frequent and impactful journals, countries, research institutes, and their relationship at social and conceptual levels.

Overall the information reported in this study could be useful to the researchers in locating the topics that need more scientific efforts, giving information to help further develop the already thriving growing field of dog cognition and behavior, thus fostering future interdisciplinary collaborations.

We used bibliometrix, a new R-tool for comprehensive science mapping analysis (Aria and Cuccurullo 2017 ), which provides various options for importing bibliographic data from scientific databases and performing bibliometrics analysis related to different items. We employed bibliometrix to analyze the sources, countries, and affiliations. It allowed us to define the structure of the topic at the conceptual level based on the co-occurrence network and thematic maps and social structure as gathered by collaboration network and world maps.

Selection strategy

Our investigation followed the Preferred Reporting Items for Systematic Reviews and Meta-Analysis (PRISMA) guidelines, illustrating the outcomes of the literature searches and article selection process (Liberati et al. 2009 ). PRISMA consists of a checklist describing the protocol adopted for selecting the collection of articles used in a systematic literature review. It is used to ensure that the selection process is replicable and transparent. We performed a computerized bibliometric analysis from January 1985 to December 2018 for articles retrieved from the Web of Science (WoS) database, which is now maintained by Clarivate Analytics, and also retrieved articles from the Science Citation Index Expanded (SCI expanded) and the Social Science Citation Index (SSCI). Data were collected in January 2020.

To identify all publications related to this field, we defined the following query: (((TS = (((dog OR dogs) AND *cogniti*) OR (canis AND familiaris AND *cogniti*))) OR (TS = ((( dog OR dogs) AND communicat*) OR (canis AND familiaris AND communicat*))) OR (TS = ((( dog OR dogs) AND behav*) OR (canis AND familiaris AND behav*))))). TS stands for topic, that is, the search of the mentioned words in the title, abstract, and keyword lists. This query was formulated after some exploratory trials in which, after using the word behav* and cogniti*, we noted that some of the publications in our personal database related to dogs’ communication did not appear. Thus, for a more comprehensive research, we also added the word communicat*. In our search, we selected original articles in the English language, including experiments (i.e., review articles and proceedings were excluded).

The information about the retrieved articles by WoS in Bib TeX format was exported into Microsoft Excel 2017. The selection involved two selectors, which reached a satisfactory agreement level (Cohen’s K  = 0.91). The choices that did not match were resolved involving a third independent researcher and the final decision was taken by the concensus among researchers (Cuccurullo et al. 2016 ).

The inclusion criteria concerning cognitive and behavioral sciences are reported in Table 1 .

Data loading and converting

Numerous software tools support science mapping analysis; however, many of these do not assist scholars in a complete recommended workflow. The most relevant tools are bibliometrix (Aria and Cuccurullo 2017 ), CitNetExplorer (van Eck and Waltman 2014 ), VOSviewer (van Eck and Waltman 2010 ), SciMAT (Cobo et al. 2012 ), and CiteSpace (Chen 2006 ). Starting from our final collection, we loaded the data (i.e., the selected papers matching the inclusion criteria, including all their metadata) and converted it into R data frame using bibliometrix (Aria and Cuccurullo 2017 ) since it contains a more extensive set of techniques and it is suitable for practitioners through Biblioshiny (Moral-Muñoz et al. 2020 ).

To investigate the interest in dog’s cognitive and behavioral sciences, the annual trend of publications from 1985 to 2018 was compared with the whole literature in the field of cognitive and behavioral studies published in the subject category “Behavioral Science,” since most of the papers related to cognition and behavior fall in this subject category. Indeed, it includes 53 journals, some of them with the main focus on cognition, such as “ Animal Cognition ” and other focusing mainly on behavior, such as “ Animal Behavior ”. However, all journals in the subject category “Behavioral Science” tend to accept studies on cognition and/or behavior.

From 2005, the studies on dogs diverged upward from the general growing trend of the papers on cognitive and behavioral studies published in the subject category “Behavioral Science” (see results). Thus, we chose to use this point to perform the following separate analysis on dogs a posteriori. A “recent analysis”, including the last 13 years (2006–2018), was used to underline emerging aspects, and an “earlier analysis”, counting the same number of years (1993–2005), was used for comparative purposes. In this way, we were able to compare the period in which there was an increase in the scientific production on dogs exceeding the trend of studies in the subject category “Behavioral Sciences” and an equivalent number of years in which the growing trend of dogs’ cognitive and behavioral studies paralleled that of behavioral sciences. This choice allowed us to test whether the exceeding trend of dog studies was simply due to an increased effort in the same topics or new topics contributed to this increase.

We analyzed the article collection using different aggregation levels. Regarding journals, bibliometrix provides many indicators, such as the number of publications, h-index (Hirsch 2005 ), g-index (Egghe 2006 ), m-index (von Bohlen und Halbach 2011 ), and the total number of citations. Thus, we reduced the variables by applying a principal component analysis with orthomax rotation, through a statistical tool for Excel (XLSTAT 2019, Addinsoft Inc.).

Co-occurrence network, collaboration network, thematic maps, and world maps are also provided. A network is a graphical representation of item co-occurrences in a set of documents. In a co-occurrence network, the items consist of terms extracted from the article keyword lists, from the titles, or from the abstracts; while in a collaboration network, the items consist of the co-authors, the author’s affiliations, or the author’s countries. A thematic map is a Cartesian representation of the term clusters identified performing a cluster analysis on a co-occurrence network. It allows for easier interpretation of the research themes developed in a framework. Finally, a world map is a geographical representation of the collaboration network of an author’s country. The analyses were based on KeyWords Plus, which are the words or phrases that frequently appear in the titles of the references cited in an article but do not appear in the title of the article itself. They are extracted from the papers using a statistical algorithm, based on the cited references in the article. This process is unique to Clarivate Analytics databases. The algorithm is based on a supervised machine learning approach that automatically assigns a set of keywords, namely, Keyword Plus, from a glossary defined by a team of experts. This approach uses the article’s bibliography to identify the research topics and then label the document with a set of Keyword Plus. The use of the KeyWords Plus offers several advantages over other databases and author’s keyword list, in such a way that the terms are extracted from a standardized glossary, defined for subject categories analyzed. It also covers a larger knowledge base and unbiased concerning the author’s subjectivity when providing keywords for their articles (Zhang et al. 2016 ). Moreover, a comparison between Keywords Plus and Author Keywords performed at the scientific and the document levels yields more Keywords Plus terms than Author Keywords, and it is more descriptive (Zhang et al. 2016 ).

Based on Keywords Plus, we obtained the co-occurrence network, which identifies the relationship between the keywords. Each keyword represents a node, or vertex, of the network, and the edge connecting two nodes is proportional to the number of times two keywords are included in the same keyword list. Stronger is an edge, higher is the relationship between two keywords within a paper (Tijssen and Van Raan 1994 ), thus allowing to provide a graphic visualization of potential relationships among keywords. In the network, it is possible to identify groups of strongly interconnected terms, which represent themes or topics. Although different algorithms exist to identify these groups, this study used the Louvain community detection algorithm (Blondel et al. 2008 ) because it gave the best results when applied to different benchmarks on Community Detection methods (Lancichinetti and Fortunato 2009 ).

The clusters identified by the co-occurrence network were plotted in a thematic map according to Callon’s centrality and Callon’s density rank values along the two axes (Callon et al. 1991 ).

The X-axis represents the centrality, that is, the degree of interaction of a network cluster in comparison with other clusters appearing in the same graph. It can be read as a measure of the importance of a theme in the development of the research field. The Y-axis symbolizes the density, which measures the internal strength of a cluster network, and it can be assumed as a measure of the theme’s development (Cahlik 2000 ; Cobo et al. 2011 , 2015 ). According to these authors, the graphical representation of themes on the four quadrants in which they are plotted allows identification of the following proprieties: (1) Motor themes (first quadrant): the cluster network is characterized by high centrality and high density, meaning that they are well developed and important for the structuring of a research field; (2) Highly developed and isolated themes (second quadrant): they are characterized by high density and low centrality, meaning that they are of limited importance for the field since they do not share important external links with other themes; (3) Emerging or declining themes (third quadrant): they have low centrality and low density, meaning that they are weakly developed and marginal. The identification of emerging or declining trends of a theme requires a longitudinal analysis, through a thematic evolution (Aria and Cuccurullo 2017 ): splitting the timespan into different timeslices allow to identify the trajectory, whereby a direction toward the top of the map over time identifies an emerging trend while a direction toward the lower left quadrant would identify a declining trend; (4) Basic and transversal themes (fourth quadrant): they are characterized by high centrality and low density, namely, they are important concerning general topics that are transversal to different research areas of the field.

The scientific collaboration analysis was used to identify the social structure of the field, through the application of the social network analysis (Newman 2001 ), applying it at an aggregated level (i.e., countries).

The comparison of the number of publications from selected papers for dogs with the general trend of all papers published in the subject category “Behavioral Science” showed that starting from 2005, there has been a sharp increase in scientific production on dogs (Fig.  1 ).

Comparative view of the annual scientific production related to cognitive studies on dogs (black line) and the trend of publication rate of research in the subject category “Behavioral Sciences” (gray line; for an obvious comparison, the line has been lowered 100 folds). It is evident that increasing trend in research on dogs starts from 2005

Data related to the main information on dogs are reported in Table 2 .

After our selection, 218 papers related to studies on dogs were retrieved in the “earlier analysis” (1993–2005), while they were 1307 in the “recent analysis” (2006–2018). This means that the scientific production on dog cognitive and behavioral studies increased sixfold. A similar finding was also observed for the Keywords Plus, Authors’ Keywords, Authors, Author Appearances, Authors of multi-authored documents, and single-authored documents. However, these data support the view that there is a considerable increase in the researchers working on the cognitive and behavioral aspects of dogs. It also appears that the contribution of a single researcher who co-authored remains almost unchanged, which means that the research effort by each researcher has not generally increased over time.

Sources impact

Dog cognitive and behavioral studies appeared in 34 different sources in “earlier analysis”, while they substantially increased in “recent analysis”, totaling to 85. The principal component analysis (PCA), carried out on the number of publications, h-index, g-index, m-index, and the total number of citations, highlighted a single principal component explaining 98.56% of the variability (Eigenvalue = 4.928, χ 2  = 584.479, P  < 0.001), with KMO  = 0.786 ensuring the sampling adequacy in the “earlier analysis”. In the “recent analysis”, the PCA detected a single component explaining most of the variability (93.709%, Eigenvalue = 4.685, χ 2  = 875.361, P  < 0.001, KMO =  0.838).

The highest score for the cognitive and behavioral sciences of dogs was the Applied Animal Behavior Science , both in “earlier analysis” and “recent analysis”. Journal of the American Veterinary Medical Association , occupying the second place in “earlier analysis”, was less utilized in “recent analysis”, not appearing in the first ten sources anymore. A lowering in score from “earlier analysis” to “recent analysis” was also observed for the third journal in the list, which was the Journal of Comparative Psychology . In “recent analysis”, Animal Cognition and Journal of Veterinary Behavior-Clinical Applications and Research both showed an increase in the score, occupying the second and third place, respectively. It is noteworthy that PLoS One acquired a high score in dog cognitive and behavioral studies in “recent analysis”. Indeed, it was not present in “earlier analysis” since it was launched in 2006, which coincides with the start of our “earlier analysis” analysis. Similar reasoning could be applied for Scientific Reports , which was launched in 2011 and is in the first ten sources in our collection related to the “recent analysis”. Full data for the sources are reported in Online Resources 1 and 2.

Country productivity and affiliations

According to our collection of the metadata, in the “earlier analysis”, 22 countries contributed to dog cognitive and behavioral studies, whereas in the “recent analysis”, they have almost been doubled (42).

Considering the number of publications related to corresponding authors, the leading countries were the USA, the UK, and Hungary in the “earlier analysis”, and these were also among the most productive in “recent analysis”. Ireland appeared as the fifth most productive country in “earlier analysis” but was much less involved in dog cognitive and behavioral studies in “recent analysis”, where it ranked at the 24th position. A slightly less engagement on the part of the Netherlands affected its ranking, pushing it down from the ten to the fifteenth position. On the contrary, Italy, contributed less in “earlier analysis”, while it appeared more productive in “recent analysis”, ranking in the third position. A similar observation holds for Austria, which was not listed in “earlier analysis”, but appeared in the first ten most productive countries in the “recent analysis”. Japan was not in the top ten contributing countries in “earlier analysis” but was indeed listed in “recent analysis”. However, this country has only earned two positions, thus maintaining its almost unchanged status in terms of its contribution. The whole data of countries’ productivity are given in Online Resources 3 and 4.

Concerning the affiliations, the Eötvös Loránd University of Hungary provided the highest contribution to the development of the dog cognitive and behavioral sciences in the “earlier analysis”, followed by the University of Toronto and the Utrecht University (the Netherlands). This ranking substantially changed in the “recent analysis”. Besides Eötvös Loránd University of Hungary, which has always been the major contributor to developing dog cognitive and behavioral sciences, all affiliations in the top ten were new, with the University of Vienna and the University of Milan occupying the second and third place, respectively, in the list. The list of the most productive affiliations can be found in Online Resources 5 and 6.

Conceptual structure

The analysis of KeyWords Plus in the “earlier analysis” identified six clusters, represented in a thematic map, according to their centrality and density ranking (Fig.  2 ). A cluster was characterized by high centrality and high density and was positioned in the first quadrant as a motor theme. It included words “beta-amyloid accumulation,” “dysfunction,” and “canine” as the most recurrent terms. A second cluster with good centrality and density identified another well-developed theme, including the KeyWords Plus “rats,” “plasma-cortisol,” and “stress” as the most co-occurring words. Another cluster was in the fourth quadrant, characterized by high centrality and a lower density with respect to the two previous clusters. It was a basic and transversal well-developed theme. The most frequent words were “comprehension,” “search behavior” “chimpanzees pan-troglodytes,” “evolution,” and “gaze.” Two other clusters were characterized by low centrality and low density, which meant they were weakly developed and marginal, and were positioned in the third quadrant. One cluster included the KeyWords Plus “animals,” “humans,” and “temperament”; the other contained, as most frequent KeyWords Plus, “classification,” “breed,” and “sex.” Finally, an isolated theme, with a high density and low centrality, including the only word “information,” was positioned in the second quadrant, which meant that it had limited importance for the field.

Thematic map showing clusters and the KeyWords Plus from 1993 to 2005 (“earlier analysis”) identified by the co-occurrence network. The X axis represents the centrality (i.e. the degree of interaction of a network cluster in comparison with other clusters) and gives information about the importance of a theme. The Y axis symbolizes the density (i.e. measures the internal strength of a cluster network, and it can be assumed as a measure of the theme’s development). Accordingly, the first quadrant identifies motor themes (i.e. well developed and important themes for the structuring of a research field); in the second quadrant are plotted highly developed and isolated themes (i.e. themes of limited importance for the field); the third quadrant contains emerging or declining themes (i.e. themes weakly developed and marginal); in the fourth quadrant falls basic and transversal themes (i.e. they concerns general topics that are transversal to different research areas of the field)

For the period characterizing the “recent analysis”, bibliometrix individuated four clusters, two in the first quadrant and two in the third (Fig.  3 ). A cluster in the first quadrant was characterized by high centrality and high density and included motor themes in dog cognitive and behavioral studies. The most occurring KeyWords Plus was “temperament,” followed by “personality” and “traits.” The second cluster in the first quadrant had a lower centrality and lower density with respect to the previous cluster and included “humans,” “performance,” and “wolves,” as the most occurring KeyWords Plus. The two clusters in the third quadrant were characterized by low centrality and low density, which meant that they were weakly developed marginal themes. A cluster included “brain,” “age,” “model,” and “social-behavior,” as most common KeyWords Plus, while the other cluster included “stress,” “welfare,” and “attachment.”

Thematic map showing clusters and the KeyWords Plus from 2006 to 2018 (“recent analysis”) identified by the co-occurrence network. The X axis represents the centrality (i.e. the degree of interaction of a network cluster in comparison with other clusters) and gives information about the importance of a theme. The Y axis symbolizes the density (i.e. measures the internal strength of a cluster network, and it can be assumed as a measure of the theme’s development). Accordingly, the first quadrant identifies Motor themes (i.e. they are well developed and important for the structuring of a research field); in the second quadrant tare plotted highly developed and isolated themes (i.e. they are of limited importance for the field); the third quadrant contains emerging or declining themes (i.e. they are weakly developed and marginal); the fourth quadrant includes basic and transversal themes (i.e. themes concerning general topics transversal to different research areas of the field)

Social structure

The USA and Canada shared the highest number of publications in “earlier analysis”, whereas other countries showed a lower rate of collaboration. In contrast, in the “recent analysis”, a radical increase in the collaboration network is evident (compare Figs.  4 and 5 ). The USA was again the leading country, sharing most publications with Canada and Australia. Hungary and Austria were at the third position, with several publications in common. It is noteworthy to underlines the radical increase in the collaboration network of the UK. The list of the most collaborative countries can be found in Online Resources 7 and 8.

World map showing research collaborations among countries from 1993 to 2005 (“earlier analysis”). Brighter blue color indicates a higher collaboration rate. Countries with less than three shared papers are not shown by connectors

World map showing research collaborations among countries from 2006 to 2018 (“recent analysis”). Brighter blue color indicates a higher collaboration rate. Countries with less than three shared papers are not shown by connectors

The first goal of the present paper was to verify whether the dog cognitive and behavioral studies were attracting the interest of scholars more than the studies on cognitive and behavioral sciences in general. It is clear from our data that scientific production for dog cognitive and behavioral studies is substantially more consistent in “recent analysis”, as compared with “earlier analysis”. Furthermore, the growing trend in studies on dogs, starting from 2005, largely exceeds the number of papers in the subject category “Behavioral Sciences,” which we have taken as a reference point for cognitive and behavioral studies in general. Our data provide scientific evidence that in recent times the interest toward the cognitive and behavioral studies on dogs is growing more than the general trend in cognitive and behavioral sciences.

Why the number of studies started to increase around 2005 is difficult to say. The most cited papers in this field came around this period and dealt with dog’ domestication (Hare et al. 2002 ; Miklósi et al. 2003 ; Hare and Tomasello 2005 ). These studies might have provided an important stimulus in raising the interest in the cognitive and behavioral studies on dogs. However, our data show that the interest on the theme related to the dog evolution remained almost unchanged from “earlier analysis” to “recent analysis”; while the studies related to temperament increased considerably (compare Figs.  2 and 3 ).

Although the total number of papers on dog cognitive and behavioral studies associated with an author’s name has increased, the contribution of a single researcher co-authoring these studies remains almost unchanged, which is in line with the general trend of scientific studies (Fanelli and Larivière 2016 ). In other words, the number of co-authors per publication increased, but the publication rate per scholar engaged in dog cognitive and behavioral studies did not. This result reflects an increasing trend in the collaboration network from the “earlier analysis” to “recent analysis” (compare Figs.  4 and 5 ).

Our second goal aimed to understand whether new research themes and/or changes in research themes accompanied the growth of scientific production in dog cognition and behavior starting from 2005. Looking at KeyWords Plus a motor theme in “earlier analysis” was characterized by studies related to aging. Studies in this field used dogs as a model to receive information about the decline of the human brain in elderly people (Bosch et al. 2011 ). This research theme received limited interest in the “recent analysis” as it showed a decrease in the centrality and density in relationship to the other themes. This means that dogs are less used as models of study for human disease, in comparison with themes aimed to study dogs per se. A similar trend was observed for the research theme related to welfare, which was characterized by good centrality and good density in “earlier analysis”, but it appeared less developed in “recent analysis” in comparison with other themes. Some of the themes maintained a high interest for both the studied periods. Indeed, the theme related to species comparison (evolution) of cognitive and behavioral skills appeared as clusters located almost at the same position in the graph. What deserved more importance from “earlier analysis” to “recent analysis” was the studies related to temperament. This theme appeared as a weakly developed and marginal theme in “earlier analysis”, while it became a motor theme in “recent analysis”. Therefore, this theme was an emerging theme in “earlier analysis”. This was a trend that had already been observed ten years before in another study (Kubinyi et al. 2009 ). Lastly, a theme including words related to breed, gender, and aggressive behavior classification, appeared as a weakly developed and marginal theme in “earlier analysis”, but it disappeared in “recent analysis”, indicating that it was already a declining theme in “earlier analysis”. It should be underlined that because of the general increase in the cognitive and behavioral studies on dogs in the “recent analysis”, all themes received an increased research effort in recent times but not with the same intensity. Thus, indicating a theme as declining should be interpreted in comparison with other themes in the same period. In this sense, the themes related to aging and welfare were less considered, thus requiring more scientific attention. Overall, both changes in research themes and new research themes contributed to the increased scientific production in dog cognitive and behavioral studies.

About the sources, Applied Animal Behavior Science received the highest score by PCA, both in “earlier analysis” and “recent analysis”. However, there was a substantial turnover for other journals, such as Journal of the American Veterinary Medical Association , which showed a high score in “earlier analysis”, but not in “earlier analysis”. On the contrary, some sources such as Animal Cognition and Journal of Veterinary Behavior-Clinical Applications and Research showed a growing trend. Besides, the new sources appearing only in “recent analysis”, such as PLoS One and Scientific Reports , caused a change in journal ranking, which also reflected as changes in topics (see below). Noteworthy, the ranking is not indicative of the value of a journal since our data do not allow to test for the quality of research reported. Moreover, our analysis refers only a limited field of studies reported in the journals, namely, studies in the cognitive and behavioral science of dogs. Other factors could also affect the ranking, such as the rejection rate, which is different between journals. In recent times, the choice for journal also depends on whether it is open access with a rapid turnover and a larger “reach”. This could be the case with more general journals, like PLoS One and Scientific Reports .

Among countries, the USA, the UK, and Hungary were the major contributors to the development in dog cognitive and behavioral studies, both in “earlier analysis” and “recent analysis”. While some countries, e.g., Ireland, showed a decline, other countries played a major role, as in the case of Italy and Austria. It should be emphasized that the numbers of research groups differ in the countries. For example, in Hungary and Austria, a single research group is responsible for almost all publications, while several groups in the US and UK contribute to the countries' achievements. The multiple-country publications (MCP), defined as the number of articles including at least a co-author working in a different country with respect to the corresponding author, showed that different countries demonstrated high levels of collaboration activity in “recent analysis”. Indeed, the inter-country collaboration appeared rich in terms of institutions forming large groups of research worldwide. On the other hand, the collaboration net was poorly developed in “earlier analysis”. This outcome is expected since the advances in technology, and the rapid processing and dissemination of scientific information have reduced the barriers of geographic distance and broadened interdisciplinary collaboration.

The data for countries were partially coherent with institutions. Indeed, the Eötvös Loránd University of Hungary provided the maximum contribution to the development of dog cognitive and behavioral sciences, both in “earlier analysis” and “recent analysis”, and the University of Vienna and the University of Milan contributed much in “recent analysis”. Regarding the journals, our analysis does not provide information about the value of a country or an institution and reflects only the studies focused on the cognitive and behavioral science of dogs. It should be remembered that the factors other than the interest in studies on the cognitive and behavioral science of dogs, can affect these outcomes. For instance, the important countries could be driven by research funding, type of institution, or animal ethics regulations. The total number of researchers can affect the number of studies regarding the cognitive and behavioral science of dogs in a country. Some institutions could have had a limited period of activity or started studies on dog cognition and behavior only a while ago. In any case, according to our goal, the present analysis offers a clear picture regarding the most productive countries and affiliations, which is useful to locate where the scientific production on cognition and behavior in dogs is more intense. Moreover, we also provide a map of collaboration, which is useful for several reasons. The ability to debate and share the experience is essential for academic and scientific accomplishment. Constructively, challenging the accepted opinions and ideas is central to their development, and international collaborations help to facilitate this. The collaboration provided a means to professional advancement and increased knowledge (Lukkonen et al. 1992 ). Beaver and Rosen ( 1979 ) concluded that scientific collaboration represents a response to the increasing professionalization of science. Persson et al. ( 2004 ) show that citations to articles resulting from international collaborations grow faster than those referring to domestic collaborations. Narin ( 1991 ) shows that internationally co-authored articles are more highly cited.

This study has several limitations, which are mainly related to the instrument of bibliometric analysis per second. Indeed, although our selection has limited false-positive, false-negative results are always present in bibliometric research, as it is impossible to generate a perfect and all-encompassing research query.

We included articles only from Web of Science (WoS), and therefore our research cannot cover the entire literature on cognition and behavior in dogs. However, it should be underlined that no scientific database is comprehensive, and each of them, including Scopus or PubMed, has its own strengths and weaknesses (Falagas et al. 2008 ). Moreover, WoS does not allow electronic access to articles published before 1985. Additionally, many other articles might have been published in not-yet-indexed journals, and therefore they will not be found in any database.

Taking all these limitations into consideration, the number of publications analyzed in this study might not precisely reflect the worldwide research activity on dog cognitive and behavioral studies, but the data presented may still provide significant insight into the evolving trends before and after the increase in dog cognitive and behavioral studies.

In conclusion, it was noted that most of the extant literature on the growing trend in the field of dog cognition and behavioral studies was published after 2000 (Arden et al. 2016 ), which is in line with our results. Indeed, an increasing trend of publications on dog cognition and behavior is evident in our analysis, which is starting to overlap the cognitive and behavioral studies on other species since 2005. Therefore, it is evident from our data that dogs are attracting the interest of scholars much more than before, and that this effect does not involve, for the same extension, the trend of the cognitive and behavioral research in general. Accordingly, the number of countries and researchers involved in these studies has also increased. In conclusion, it seems that in addition to being the man’s best friend (Miklósi and Topál 2013 ), the dog is also becoming the scholar’s best friend.

Data availability

The datasets generated and/or analyzed during the current study are available from the corresponding author on request.

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Online Resource 1

Information about sources ordered according to the scores (S) of the principal component analysis from 1993 to 2005 ("earlier analysis"). NP: the number of publications; TC: total citations; h-index: journal’s number of published articles (h), each of which has been cited in other papers at least h time; g-index: the largest number such that the top g articles received at least g 2 citations; m-index: the ratio h/n, where h is the h-index and n the number of years since the first published paper. The first three items are in bold (XLSX 19 KB)

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Information about sources ordered according to the scores (S) of the principal component analysis from 2006 to 2018 ("recent analysis": RA). NP: the number of publications; TC: total citations; h-index: journal’s number of published articles (h), each of which has been cited in other papers at least h time; g-index: the largest number such that the top g articles received at least g 2 citations; m-index: the ratio h/n, where h is the h-index and n the number of years since the first published paper. The first three items are in bold (XLSX 25 KB)

Online Resource 3

Information about countries production ordered by the total number of publications by corresponding authors (CNP) from 1993 to 2005 ("earlier analysis"). CFreq: citation frequency according to the publication by the corresponding author; SCP: single country publications (i.e. number of articles in which all authors belong to the same country); MCP: multiple countries publications (i.e. number of articles including at least a co-author working in a different country with respect to the corresponding author); TNP: the number of publications; TC: total citation received; The first three items are in bold (XLSX 11 KB)

Online Resource 4

Information about countries production ordered by the total number of publications by corresponding authors (CNP) from 2006 to 2018 ("recent analysis": RA). CFreq: citation frequency according to the publication by the corresponding author; SCP: single-country publications (i.e. number of articles in which all authors belong to the same country); MCP: multiple-country publications (i.e. number of articles including at least a co-author working in a different country with respect to the corresponding author); TNP: number of publications; TC: total citation received; The first three items are in bold (XLSX 13 KB)

Online Resource 5

The most productive affiliations from 1993 to 2005 ("earlier analysis": EA). The first three items are in bold (XLSX 11 KB)

Online Resource 6

The most productive affiliations from 2006 to 2018 ("recent analysis": RA). The first three items are in bold (XLSX 16 KB)

Online Resource 7

Numbers of shared publications between countries in the period 1993–2005 ("earlier analysis": EA) (XLSX 9 KB)

Online Resource 8

Numbers of shared publications between countries in the period 2006–2018 ("recent analysis": RA) (XLSX 9 KB)

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Aria, M., Alterisio, A., Scandurra, A. et al. The scholar’s best friend: research trends in dog cognitive and behavioral studies. Anim Cogn 24 , 541–553 (2021). https://doi.org/10.1007/s10071-020-01448-2

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Some dog breeds are better at learning by observing humans than others

Have you ever noticed how different dog breeds seem to be better at social learning, from observing humans, than others?

A recent study from the Department of Ethology at Eötvös Loránd University has shed some intriguing light on this subject. Their research findings suggest that some breeds of dogs may have more of an edge when it comes to gleaning insights from observing human actions.

A question that often pops up when debating dog behavior is whether the breed of the dog has any influence on the results of such studies. And, if so, why or why not?

There are hundreds of distinct dog breeds out there. Each breed varies not just in their looks, but also in their behaviors. It's quite common to assume that a terrier might act differently than a Husky. Or, similarly, that a Border Collie might exhibit different tendencies than a pointer.

Differences in social learning ability across dog breeds

Ethologists have indeed found significant breed differences in many experiments. However, when it came to social learning, breed difference seemed a bit elusive. Social learning is the ability to learn by watching others' actions.

Over the last two decades, regardless of breed, dogs have shown an uncanny ability to learn from other dogs. Also, and even more impressively, they can also learn from watching humans.

But this new study, published in the journal 'Animals', has brought a fresh perspective. The researchers have shown that certain dog breeds might have a sharper inclination towards human behavior.

This tendency can be a real game-changer when dogs face a tricky spatial problem-solving task. An example of this would be finding a detour around an obstacle.

How the new social learning study was conducted

The researchers behind this compelling study are Dr. Péter Pongrácz, a pioneer in social learning experiments with dogs. Also, his partner, Petra Dobos, an undergraduate student with tremendous potential.

Dobos's bachelor thesis aimed to unearth breed-related differences that could explain variations in dogs' social learning skills.

The dynamic duo employed a novel approach by grouping dog breeds into two functional groups. First were the cooperative working dogs. These include herding dogs, pointers, and retrievers who work closely with their human handlers.

Second are independent working dogs. This group includes terriers, sighthounds, spitz type breeds and livestock guarding dogs. They function mostly without continuous human guidance.

Testing dog breeds with the 'V-shaped fence detour' test

In the study, nearly 100 dogs were subjected to the renowned 'V-shaped fence detour' test . This task involves the dog finding a reward placed behind a transparent wire-mesh fence. This is a feat that many dogs struggle to accomplish within the standard one-minute trial duration.

Each dog underwent three consecutive trials. Each trial had a control group receiving no additional help.

The other group, however, got a leg up by observing humans. Petra walked around the fence before the second and third trials. This demonstrated the correct path to the reward.

As they dove into the data, the researchers found that both independent and cooperative breeds had a similar success rate in the control condition. This was to be expected considering the inherent difficulty of the detour task for dogs.

What the researchers discovered

Interestingly, when the dogs were given the chance to observe a human demonstrator navigate the detour, the cooperative working dog breeds clearly had the upper hand.

Unlike the independent dogs, cooperative dogs significantly improved their detour time compared to their initial trial, even though both groups observed the same human demonstration.

Dr. Pongrácz, the supervisor of the research program, reflected on the study's findings. "The beauty of this finding lays in the fact that we did not find any specific dog breed to be especially talented in social learning," he said.

The groups consisted of unrelated dog breeds, positioned at distant branches on the genetic ‘tree' of dogs. This highlights the role of functional breed selection that possibly focuses on enhancing dogs' attention and interest towards human behavior.

More research planned on dog breeds and social learning

The research duo has plans to delve deeper into these breed differences in future studies. Petra voiced the team's ambition.

"We won't stop here of course. There are so many things still to discover. For example, whether the cooperative and independent dogs would equally regard our attention calling words interesting. Or, would we find the cooperative breeds as being more attracted to human encouragement?"

In conclusion, this study has taken a significant stride in understanding the nuanced ways in which breed differences may influence a dog's ability to learn from human actions. It paves the way for more fascinating discoveries in the world of ethology.

More about relationship between dogs and humans

Dogs and humans have shared a unique bond for thousands of years. These bonds have spanned different cultures, regions, and societies.

This human/dog relationship is marked by mutual companionship, assistance, and love. In fact, dogs were the first species to be domesticated by humans. This occured somewhere between 20,000 and 40,000 years ago. Here's a deeper look into this special connection:

Domestication

Wolves, the ancestors of all modern dogs, were likely first attracted to human settlements due to the availability of food scraps. Over time, wolves that were more tolerant of humans and human behavior likely had an advantage. This led to a process of self-domestication.

Humans, in turn, found value in these animals. Dogs were used not only as sources of food and fur, but also as guards, hunters, and companions.

Co-Evolution

Humans and dogs have co-evolved. This means that as humans have shaped dogs through selective breeding, dogs have also influenced human evolution.

Some theories suggest that the collaboration between dogs and early humans allowed the latter to hunt more efficiently and have a more regular source of nutrients. These nutrients might have contributed to the development of our large brains.

Communication

Dogs have developed a unique ability to understand human signals, gestures, and speech. They can read our body language and vocal cues. They respond to our commands. Most dogs can even interpret our emotions.

This capacity for understanding and communication far exceeds that of any other animal species. It is a testament to the deep connection between dogs and humans.

Emotional Bonding

Dogs and humans have the capacity to form deep emotional bonds. Dogs are known for their loyalty and are often considered members of the family.

Research has found that interacting with dogs can release the hormone oxytocin, known as the "love hormone", in both the human and the dog. Oxytocin further reinforces this human/dog bond.

Assistance Roles

Dogs play many crucial roles in human society beyond companionship. They can be trained as service dogs to assist people with disabilities, as therapy dogs to provide emotional support, and as working dogs in various professions. These include search and rescue, law enforcement, and herding livestock.

Health Benefits

Interacting with dogs has been shown to provide numerous health benefits for humans. These include lower stress levels, reduced blood pressure, increased physical activity, and improved mental health.

Some studies have also shown that children who grow up with dogs have a lower risk of developing allergies and asthma.

Cultural Significance

Dogs hold significant cultural importance in many societies. They are often featured in art, literature, and mythology. In some cultures, dogs are revered and considered sacred, while in others, they are cherished pets and family members.

The relationship between humans and dogs continues to evolve as we learn more about these remarkable animals. While the form of the relationship can vary from place to place and person to person, the mutual respect and love that form its foundation remain constant.

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Maddie's Insights - Impact of Classical Counterconditioning (Quiet Kennel Exercise) on Barking in Shelter Dogs (OnDemand)

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A major welfare concern in animal shelters is excessive barking from kenneled dogs. This contributes to noise pollution, can cause hearing damage, and has a negative impact on all animals within earshot. This study demonstrated that by implementing a simple classical counterconditioning exercise, consisting of instructing all ward passers-through to toss treats to each dog regardless of behavior exhibited, could change the emotional state of dogs from negative to more positive, thus reducing fear and frustration that often leads to excessive barking.

Data analysis showed that implementation of this simple exercise did reduce barking at clinically relevant levels even with a changing shelter population.  Additionally, for each percent increase in compliance rate, there was an average decrease of 17.3 dB in the maximum decibel reading. Moreover, an overall positive change in attitude from most dogs towards visitors further showed the positive effects of the study.

Learning Objectives:

1.     Attendees will learn to identify and apply basic learning theory, specifically focusing on Classical (Pavlovian) Conditioning, to dogs in a kennel setting.

2.     Attendees will learn to identify risks and consequences of noise pollution in shelters, and also ways to reduce it to improve welfare of all species in the shelter.

Presenters :  Sara L. Bennett, DVM, MS, DACVB (Behavior),  Clinical Assistant Professor, Veterinary Behavior Specialist, Department of Clinical Sciences, North Carolina State University, College of Veterinary Medicine and  Jamirelis Carrero (Jami),  DVM candidate, North Carolina State University College of Veterinary Medicine

This webinar has been pre-approved for 1.0 Certified Animal Welfare Administrator continuing education credits by The Association for Animal Welfare Advancement and by the National Animal Care & Control Association. It has also been approved for 1 hour of continuing education credit in jurisdictions which recognize RACE approval. Complete the quiz to earn continuing education credit for CAWA and NACA.

Visit Maddie's Pet Forum to comment, follow a discussion or ask questions:  https://maddies.fund/MIQuietKe...

keywords  Maddie's Insights, Sara L.Bennett, dog behavior, enrichment, handling and training, pavlovian conditioning, barking in kennel dogs, classical counterconditioning exercise, dog treats, emotional state of dogs

Sara L. Bennett, DVM, MS, DACVB

Clinical assistant professor, veterinary behavior specialist, north carolina state university, college of veterinary medicine.

Dr. Bennett received her DVM degree in 2006 from Purdue University and later completed a residency in animal behavior with a focus on shelter medicine, obtained a master of science degree, and became certified as a diplomate of the American College of Veterinary Behaviorists in 2012. She spent time in specialty private practice and held a position as co-instructor for the University of Florida College of Veterinary Medicine Shelter Medicine Program, focusing on Shelter Animal Behavior and Welfare, for 8 years. She is currently a clinical assistant professor of clinical sciences in the behavior department at the College of Veterinary Medicine at North Carolina State University. Dr. Bennett particularly enjoys addressing problem behaviors in sheltered and rescued animals through clinical practice and research, thereby helping to protect animal welfare, making the sheltering experience less stressful, strengthening the human-animal bond, and increasing the likelihood that pets’ new homes become their forever homes.

Jamirelis Carrero

Dvm candidate, north carolina state university college of veterinary medicine.

Jamirelis Carrero (Jami) is an incoming third year vet student at North Carolina State University. Born and raised in Puerto Rico, and Jami has a bachelor’s degree in animal science from the University of Puerto Rico at Mayagüez Campus. After completing a summer research study under the guidance of Dr. Sarah Bennett and the Veterinary Scholars Program, Jami is passionate about finding ways to use behavior modification techniques to improve the quality of care that animals receive in shelters and clinical scenarios.

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To optimize guide-dog robots, first listen to the visually impaired

What features does a robotic guide dog need? Ask the blind, say the authors of an award-winning paper. Led by researchers at the University of Massachusetts Amherst, a study identifying how to develop robot guide dogs with insights from guide dog users and trainers won a Best Paper Award at CHI 2024: Conference on Human Factors in Computing Systems (CHI).

Guide dogs enable remarkable autonomy and mobility for their handlers. However, only a fraction of people with visual impairments have one of these companions. The barriers include the scarcity of trained dogs, cost (which is $40,000 for training alone), allergies and other physical limitations that preclude caring for a dog.

Robots have the potential to step in where canines can't and address a truly gaping need -- if designers can get the features right.

"We're not the first ones to develop guide-dog robots," says Donghyun Kim, assistant professor in the UMass Amherst Manning College of Information and Computer Science (CICS) and one of the corresponding authors of the award-winning paper. "There are 40 years of study there, and none of these robots are actually used by end users. We tried to tackle that problem first so that, before we develop the technology, we understand how they use the animal guide dog and what technology they are waiting for."

The research team conducted semistructured interviews and observation sessions with 23 visually impaired dog-guide handlers and five trainers. Through thematic analysis, they distilled the current limitations of canine guide dogs, the traits handlers are looking for in an effective guide and considerations to make for future robotic guide dogs.

One of the more nuanced themes that came from these interviews was the delicate balance between robot autonomy and human control. "Originally, we thought that we were developing an autonomous driving car," says Kim. They envisioned that the user would tell the robot where they want to go and the robot would navigate autonomously to that location with the user in tow.

This is not the case.

The interviews revealed that handlers do not use their dog as a global navigation system. Instead, the handler controls the overall route while the dog is responsible for local obstacle avoidance. However, even this isn't a hard-and-fast rule. Dogs can also learn routes by habit and may eventually navigate a person to regular destinations without directional commands from the handler.

"When the handler trusts the dog and gives more autonomy to the dog, it's a bit delicate," says Kim. "We cannot just make a robot that is fully passive, just following the handler, or just fully autonomous, because then [the handler] feels unsafe."

The researchers hope this paper will serve as a guide, not only in Kim's lab, but for other robot developers as well. "In this paper, we also give directions on how we should develop these robots to make them actually deployable in the real world," says Hochul Hwang, first author on the paper and a doctoral candidate in Kim's robotics lab.

For instance, he says that a two-hour battery life is an important feature for commuting, which can be an hour on its own. "About 90% of the people mentioned the battery life," he says. "This is a critical part when designing hardware because the current quadruped robots don't last for two hours."

These are just a few of the findings in the paper. Others include: adding more camera orientations to help address overhead obstacles; adding audio sensors for hazards approaching from the occluded regions; understanding 'sidewalk' to convey the cue, "go straight," which means follow the street (not travel in a perfectly straight line); and helping users get on the right bus (and then find a seat as well).

The researchers say this paper is a great starting point, adding that there is even more information to unpack from their 2,000 minutes of audio and 240 minutes of video data.

Winning the Best Paper Award was a distinction that put the work in the top 1% of all papers submitted to the conference.

"The most exciting aspect of winning this award is that the research community recognizes and values our direction," says Kim. "Since we don't believe that guide dog robots will be available to individuals with visual impairments within a year, nor that we'll solve every problem, we hope this paper inspires a broad range of robotics and human-robot interaction researchers, helping our vision come to fruition sooner."

Other researchers who contributed to the paper include:

Ivan Lee, associate professor in CICS and a co-corresponding author of the article along with Donghyun, an expert in adaptive technologies and human-centered design; Joydeep Biswas, associate professor at the University of Texas Austin, who brought his experience in creating artificial intelligence (AI) algorithms that allow robots to navigate through unstructured environments; Hee Tae Jung, assistant professor at Indiana University, who brought his expertise in human factors and qualitative research to participatory study with people with chronic conditions; and Nicholas Giudice, a professor at the University of Maine who is blind and provided valuable insight and interpretation of the interviews.

Ultimately, Kim understands that robotics can do the most good when scientists remember the human element. "My Ph.D. and postdoctoral research is all about how to make these robots work better," Kim adds. "We tried to find [an application that is] practical and something meaningful for humanity."

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• Engineering
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• Neural Interfaces
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Story Source:

Materials provided by University of Massachusetts Amherst . Note: Content may be edited for style and length.

Journal Reference :

• Hochul Hwang, Hee-Tae Jung, Nicholas A Giudice, Joydeep Biswas, Sunghoon Ivan Lee, Donghyun Kim. Towards Robotic Companions: Understanding Handler-Guide Dog Interactions for Informed Guide Dog Robot Design . CHI 2024: Conference on Human Factors in Computing Systems , 2024 DOI: 10.1145/3613904.3642181

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• Open access
• Published: 01 October 2021

Canine hyperactivity, impulsivity, and inattention share similar demographic risk factors and behavioural comorbidities with human ADHD

• Sini Sulkama 1 , 2 , 3   na1 ,
• Jenni Puurunen 1 , 2 , 3   na1 ,
• Milla Salonen   ORCID: orcid.org/0000-0002-9901-7923 1 , 2 , 3 ,
• Salla Mikkola   ORCID: orcid.org/0000-0003-0333-6207 1 , 2 , 3 ,
• Emma Hakanen   ORCID: orcid.org/0000-0002-8790-0225 1 , 2 , 3 ,
• César Araujo 1 , 2 , 3 &
• Hannes Lohi   ORCID: orcid.org/0000-0003-1087-5532 1 , 2 , 3  

Translational Psychiatry volume  11 , Article number:  501 ( 2021 ) Cite this article

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Attention-deficit hyperactivity disorder (ADHD) is a prevalent neurodevelopmental disorder impairing the quality of life of the affected individuals. The domestic dog can spontaneously manifest high hyperactivity/impulsivity and inattention which are components of human ADHD. Therefore, a better understanding of demographic, environmental and behavioural factors influencing canine hyperactivity/impulsivity and inattention could benefit both humans and dogs. We collected comprehensive behavioural survey data from over 11,000 Finnish pet dogs and quantified their level of hyperactivity/impulsivity and inattention. We performed generalised linear model analyses to identify factors associated with these behavioural traits. Our results indicated that high levels of hyperactivity/impulsivity and inattention were more common in dogs that are young, male and spend more time alone at home. Additionally, we showed several breed differences suggesting a substantial genetic basis for these traits. Furthermore, hyperactivity/impulsivity and inattention had strong comorbidities with compulsive behaviour, aggressiveness and fearfulness. Multiple of these associations have also been identified in humans, strengthening the role of the dog as an animal model for ADHD.

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Introduction.

Attention-deficit hyperactivity disorder (ADHD) is a highly heritable, childhood-onset neurodevelopmental disorder with an estimated worldwide prevalence of 2–7% in humans [ 1 , 2 ]. It is characterised by the presence of persistent and inappropriate levels of motor overactivity, impulsivity and inattention [ 1 , 3 ], caused by abnormalities in attention and reward processing, inhibitory control and emotional regulation [ 4 , 5 ]. ADHD can be classified into three different presentations: predominantly hyperactive/impulsive, predominantly inattentive or combined type [ 3 ]. Often, ADHD persists into adulthood with several comorbidities, such as learning impairments, autism spectrum disorder and anxiety disorders [ 1 , 4 ], making the disorder a detrimental condition if not diagnosed and treated appropriately.

Impulsivity, a component of human ADHD [ 3 ], is characterised by impaired motor inhibitory control and an inability to tolerate gratification delay [ 6 ]. It is a dimension of the normal personality continuum observed across species, including humans and dogs [ 6 , 7 ]. However, high levels of impulsivity are considered abnormal and are also associated with other detrimental behaviours, such as aggression, in several species, including humans [ 8 , 9 , 10 ], rodents [ 11 , 12 ] and dogs [ 13 , 14 , 15 , 16 , 17 ].

Currently, the under-recognition of ADHD hinders its management in humans [ 1 , 18 ]. This mostly results from the lack of appropriate animal models of ADHD [ 18 ]. Studies suggest that 12–15% [ 19 , 20 ] and 20% [ 20 ] of dogs naturally manifest high levels of hyperactivity/impulsivity and inattention, respectively, making the domestic dog a highly potential animal model for ADHD [ 21 , 22 , 23 ]. Moreover, these traits are mediated by the same behavioural [ 21 , 24 ], biological [ 17 , 25 ] and genetic [ 24 , 26 ] factors in both dogs and humans, and dogs also respond to medication used to treat ADHD in humans [ 27 , 28 ]. Furthermore, dogs have even more benefits over the classical animal models, rodents. During domestication, dogs were exposed to similar environmental factors and experienced convergent social evolution [ 29 ]. They are also comparable to humans in many complex social cognition tasks [ 30 ], genetics [ 31 ], body size and physiology and shared environment and lifestyle. The latter feature makes the dog a highly intriguing model for ADHD, as, despite the high heritability estimates, nongenetic factors also contribute to the aetiology of ADHD [ 32 ]. However, it is not well known which environmental factors affect ADHD outcomes and to what extent.

To obtain study cohorts truly resembling human ADHD, objective and reliable phenotyping of dogs is required. This can be achieved via owner-filled questionnaires as owners know their dogs’ behaviour well [ 33 ] and the reliability of questionnaires is usually good [ 34 , 35 ]. Based on a survey measuring ADHD in children [ 36 ], Vas and colleagues [ 21 ] developed an owner-filled dog-ADHD questionnaire, which can reliably measure hyperactivity, impulsivity and attention in dogs. This questionnaire was recently validated, and high levels of impulsivity and inattention were associated with a lower performance in a cognitive task [ 37 ].

In this study, we have utilised the same questions developed by Vas et al. [ 21 ] as a part of our comprehensive canine behavioural questionnaire. We aimed to examine the demographic, environmental and behavioural factors associated with canine hyperactivity/impulsivity and inattention in a study cohort of over 11,000 Finnish pet dogs. Identification of associated factors could more efficiently help to prevent and manage abnormal levels of hyperactivity/impulsivity and inattention in dogs and could also benefit human ADHD research.

Material and methods

Data collection, questionnaire.

To collect behavioural and background information from Finnish pet dogs, we designed an online owner-completed behavioural questionnaire. The questionnaire included questions about seven different canine behavioural traits: fear, aggressiveness, noise sensitivity, fear of surfaces and heights, hyperactivity/impulsivity and inattention, separation-related behaviour and compulsive behaviour. Additionally, the questionnaire included a large background section covering demographic and environmental questions related to the dog’s life history. Questionnaire replies were collected from February 2015 to September 2018. The questionnaire and more details about behavioural trait categorisation can be found as Supplementary material in the article by Salonen et al. [ 20 ]. This current study is based on the same data as the article by Salonen et al. [ 20 ] but here we utilised large multivariate analyses. Here, we studied the demographic and environmental factors associated with hyperactivity/impulsivity and inattention.

Hyperactivity/impulsivity and inattention

To measure individual differences in hyperactivity/impulsivity and inattention, we used the dog ADHD survey developed and validated by Vas et al. [ 21 ], which is based on a validated parent-report rating scale measure of ADHD and related problems in children (the ADHD RS Parent version [ 36 ]). The survey includes 13 statements (items) (Supplementary Table S1 ) concerning hyperactive, impulsive and inattentive behaviour, which were translated into Finnish. The dog owners were asked to answer how often the statement is true for their dog on a four-point Likert scale (1 = never, 2 = rarely, 3 = often, 4 = very often). Paralleling the original factor structure of Vas et al. [ 21 ], a principal component analysis with a promax rotation divided the questionnaire statements into two components, hyperactivity/impulsivity and inattention, which consisted of five and seven statements, respectively. One statement (Item 11: 'It is likely to react hastily and that’s why it is failing tasks') loaded equally on both components and was thus excluded from the analysis. We calculated component scores of hyperactivity/impulsivity and inattention for each dog, with higher component scores indicating a higher level of hyperactivity/impulsivity or inattention. Detailed information about the statements and components can be found in Supplementary Material in the article of Salonen et al. [ 20 ]. The Finnish translation of the questionnaire was recently validated in Salonen et al. [ 38 ].

Demographic, behavioural and environmental variables

Before statistical analyses, we edited some demographic and environmental variables derived from the behavioural questionnaire. We created some new variables already described in Puurunen et al. [ 39 ], Hakanen et al. [ 40 ] and Mikkola et al. [ 41 ]. All explanatory variables derived from the behavioural questionnaire are explained in detail in Supplementary Table S2 .

Briefly, we included 22 breeds with adequate sample size and mixed breed dogs and grouped other breeds under breed group other. A new variable 'body size' was created by assessing the average heights of breeds and categorised the dogs into small, medium and large dogs (Supplementary Methods). As body size could not be determined for mixed breed dogs, they were excluded when body size was included in the analysis. To quantify the environmental land-use in the dog’s home place, we generated a continuous variable 'urban environment score' (Supplementary Methods), with a higher urban environment score indicating a higher proportion of built environment.

Additionally, we created three categorical behavioural variables: 'compulsive behaviour', 'aggressiveness' and 'fearfulness' (Supplementary Methods). In all three traits, dogs were divided into three groups: low, moderate and high. The low group included dogs that never showed compulsive, aggressive or fearful behaviour, the moderate group included dogs that showed these behaviours no more than occasionally, and the high group included the dogs with regular compulsive, aggressive or fearful behaviour at least in one subtrait.

Statistical analyses

All statistical analyses were conducted in R version 3.6.2 [ 42 ]. Generalised linear models were used to analyse the associations between demographic, environmental and behavioural variables and hyperactivity/impulsivity and inattention. Gamma distribution with a log link function was used in both hyperactivity/impulsivity and inattention models, as it best fitted the data. The initial data consisted of 13,715 dogs in 264 breeds. After excluding individual dogs with missing or incomplete responses in the studied explanatory variables, the data included 6,400 dogs in both hyperactivity/impulsivity and inattention.

We used the hyperactivity/impulsivity and inattention component scores as continuous response variables in the analyses. Fourteen explanatory variables were selected for the analyses based on previous literature. Age, sex, sterilisation, breed and body size were included as demographic explanatory variables, weaning age, activities/training, daily exercise, owner’s dog experience, daily time spent alone and urban environment score as environmental explanatory variables, and compulsive behaviour, fearfulness and aggressiveness as behavioural explanatory variables.

We used a forward stepwise AIC (Akaike Information Criterion) model selection to find the best fitting model starting with a model including age and sex as explanatory variables. Sexes and age groups differed in prevalence in our previous study [ 20 ]. The AIC model selection and the final models are presented in Supplementary Table S3 . To maximise sample sizes, we created new subsets of the initial data after model selection by including all dogs that had missing responses in the explanatory variables that were not selected in the final models. As a result, the final datasets consisted of 11,539 and 11,164 individuals in hyperactivity/impulsivity and inattention, respectively.

Model fit was assessed carefully (Supplementary Methods). After fitting the model, estimated marginal means were calculated with the package 'emmeans' [ 43 ] to obtain the adjusted means and confidence limits for categorical explanatory variables. The package 'effects' [ 44 ] in R was used to obtain the effects of continuous explanatory variables, adjusting for other variables in the models. Analysis of variance (ANOVA) was conducted with the package 'car' [ 45 ] in R to get the overall effects of the explanatory variables on hyperactivity/impulsivity and inattention scores.

Based on previous studies, we formed several a priori contrasts between the levels of explanatory variables. We hypothesised that younger dogs are more hyperactive and impulsive and more inattentive than older dogs [ 21 , 22 , 46 , 47 , 48 ]. Besides, in both hyperactivity/impulsivity and inattention, we hypothesised that dogs participating in activities or training often (at least weekly) would differ from dogs participating in activities or training never or seldom [ 21 , 22 , 49 ]. In hyperactivity/impulsivity, we hypothesised that large dogs would differ from small dogs in their hyperactivity/impulsivity [ 21 , 48 ]. Furthermore, we hypothesised that dogs in high compulsive behaviour, high aggressiveness and high fearfulness groups would be more hyperactive and impulsive than dogs in low compulsive behaviour, low aggressiveness and low fearfulness groups, respectively [ 19 ].

The package 'emmeans' [ 43 ] in R was used to examine all pairwise comparisons between the levels of the categorical explanatory variables and to examine the a priori contrasts. We corrected p values for false discovery rate (FDR), except contrasts chosen a priori. We set the significance cut-off at p value <0.05.

Study cohort and demographics

We studied the effects of environmental, demographic and behavioural factors on hyperactivity/impulsivity and inattention in study cohorts of 11,539 and 11,164 dogs, respectively. The hyperactivity/impulsivity score varied from −1.62 to 5.23 (mean −0.01) and the inattention score from −1.81 to 4.81 (mean −0.01). In both hyperactivity/impulsivity and inattention, 51% of the dogs were females. The age of the dogs varied from 2 months to 17.9 years, with a mean of 4.7 years in both traits. More detailed demographics are presented in Supplementary Table S4 .

Factors associated with hyperactivity/impulsivity

Several demographic, environmental and behavioural variables, including age, sex, breed, body size, daily exercise, daily time spent alone, owner’s dog experience, compulsive behaviour, aggressiveness and fearfulness were associated with canine hyperactivity/impulsivity (Table 1 ).

We examined breed differences in hyperactivity/impulsivity in 23 breeds, including other groups consisting of all the other breeds in the data. We detected significant differences in hyperactivity/impulsivity scores between dog breeds. The breeds with the highest scores were Cairn Terrier, Jack Russell Terrier, German Shepherd and Staffordshire Bull Terrier. The breeds with the lowest scores were Chinese Crested Dog, Rough Collie and Chihuahua (Fig. 1A ). The largest pairwise differences were found between Chihuahua and German Shepherd Dog (z-ratio = −6.07, df = 1, p  = 0.0008), Chihuahua and Jack Russell Terrier (z-ratio = −5.41, df = 1, p  = 0.0008), Chihuahua and Staff. Bull Terrier (z-ratio = −5.35, df = 1, p  = 0.0008) and Chihuahua and other (z-ratio = −5.22, df = 1, p  = 0.0008). All pairwise breed differences are presented in the Supplementary Dataset.

The effects of breed ( A ), age ( B ), sex ( C ), compulsive behaviour ( D ), aggressiveness ( E ) and fearfulness ( F ) on canine hyperactivity/impulsivity. Grey area ( B ) and error bars ( A , C , D , E , F ) indicate 95% confidence limits. N  = 11,539.

Both the age and the sex of the dog were associated with hyperactivity/impulsivity (Table 1 , Supplementary Table S5 , and Fig. 1B , C). Hyperactivity/impulsivity scores were highest in young dogs ( F  = 295.93, df = 1, p  < 0.0001) as hypothesised a priori. Male dogs had higher hyperactivity/impulsivity scores than females (z-ratio = 8.27, df = 1, p  < 0.0001). There was also an association between body size and hyperactivity/impulsivity (Supplementary Table S5 and Supplementary Fig. S1A ). Medium-sized dogs had higher hyperactivity/impulsivity scores than small (z-ratio = 4.74, df = 1, p  = 0.0008) or large (z-ratio = 2.78, df = 1, p  = 0.0224) dogs, and, as we hypothesised in our a priori contrast, there was also a significant difference between small and large dogs (z-ratio = −2.40, df = 1, p  = 0.0166).

Dogs getting less daily exercise and spending more time alone had higher hyperactivity/impulsivity scores (Supplementary Table S5 and Supplementary Fig. S1B, C ). Dogs getting less than 1 h of exercise per day had higher hyperactivity/impulsivity scores than dogs exercising more than 3 h (z-ratio = 4.75, df = 1, p  = 0.0008), 2–3 h (z-ratio = 3.96, df = 1, p  = 0.0008), or 1–2 h (z-ratio = 2.5, df = 1, p  = 0.0416) per day. Additionally, dogs exercising 1–2 h had higher hyperactivity/impulsivity scores when compared with dogs exercising 2–3 h (z-ratio = 2.66, df = 1, p  = 0.0297) or more than 3 h (z-ratio = 3.74, df = 1, p  = 0.0014) daily. Dogs that spent alone more than 8 h daily had higher hyperactivity/impulsivity scores than dogs spending less than 3 h (z-ratio = 3.99, df = 1, p  = 0.0008), 3–6 h (z-ratio = 4.83, df = 1, p  = 0.0008) or 6–8 h (z-ratio = 3.48, df = 1, p  = 0.0031) alone. The owner’s dog experience was associated with higher hyperactivity/impulsivity scores (Supplementary Table S5 and Supplementary Fig. S1D ). If the dog was not the owner’s first dog, it was more likely to have a higher hyperactivity/impulsivity score than if the dog was the owner’s first dog (z-ratio = 4.65, df = 1, p  = 0.0001).

Compulsive, aggressive and fearful dogs were reported to have higher hyperactivity/impulsivity scores (Supplementary Table S5 and Fig. 1D , E, F). As we hypothesised a priori, dogs showing high levels of compulsive behaviour had higher hyperactivity/impulsivity scores when compared with dogs showing low levels of compulsive behaviour (z-ratio = 21.50, df = 1, p  < 0.0001). Also, dogs showing high levels of compulsive behaviour had higher hyperactivity/impulsivity scores than dogs showing moderate levels of compulsive behaviour (z-ratio = 13.78, df = 1, p  = 0.0008), and dogs showing moderate levels of compulsive behaviour had higher hyperactivity/impulsivity scores than dogs showing low levels of compulsive behaviour (z-ratio = 11.61, df = 1, p  = 0.0008). Similarly, as we hypothesised a priori, dogs showing high levels of aggressiveness had higher hyperactivity/impulsivity scores when compared with dogs showing low levels of aggressiveness (z-ratio = 10.21, df = 1, p  < 0.0001). Dogs showing high levels of aggressiveness had also higher hyperactivity/impulsivity scores than dogs showing moderate levels of aggressiveness (z-ratio = 4.71, df = 1, p  = 0.0008), and dogs showing moderate levels of aggressiveness had higher hyperactivity/impulsivity scores than dogs showing low levels of aggressiveness (z-ratio = 6.80, df = 1, p  = 0.0008). Furthermore, as also hypothesised a priori, dogs showing high levels of fearfulness had higher hyperactivity/impulsivity scores when compared with dogs showing low levels of fearfulness (z-ratio = 20.01, df = 1, p  < 0.0001). Dogs showing high levels of fearfulness had also higher hyperactivity/impulsivity scores than dogs showing moderate levels of fearfulness (z-ratio = 10.93, df = 1, p  = 0.0008), and dogs showing moderate levels of fearfulness had higher hyperactivity/impulsivity scores than dogs showing low levels of fearfulness (z-ratio = 8.77, df = 1, p  = 0.0008).

Factors associated with inattention

Statistical analysis identified several demographic, environmental and behavioural factors associated with inattention scores, including breed, age, sex, daily time spent alone, activities/training, compulsive behaviour, aggressiveness and fearfulness (Table 1 ).

We detected significant differences in inattention scores between dog breeds. The breeds with the highest scores were Cairn Terrier, Golden Retriever and Finnish Lapponian Dog. The breeds with lowest scores were Spanish Water Dog, Miniature Poodle and Border Collie (Fig. 2A ). The largest pairwise differences were found between Border Collie and other (z-ratio = −8.01, df = 1, p  = 0.0005), Border Collie and Finnish Lapponian Dog (z-ratio = −7.60, df = 1, p  = 0.0005), Border Collie and mixed breed (z-ratio = −6.83, df = 1, p  = 0.0005) and Border Collie and Wheaten Terrier (z-ratio = −6.54, df = 1, p  = 0.0005). All pairwise breed differences are presented in the Supplementary Dataset.

The effects of breed ( A ), sex ( B ), compulsive behaviour ( C ), aggressiveness ( D ) and fearfulness ( E ) on canine inattention. Error bars indicate 95% confidence limits. N  = 11,164.

Age of the dog was associated with inattention (Table 1 and Supplementary Fig. S2A ). Inattention scores were highest in young dogs (linear effect: F  = 8.73, df = 1, p  = 0.0031, quadratic effect: F  = 0.56, df = 1, p  = 0.561) as we hypothesised a priori. Significant differences were also found between sexes as males had higher inattention scores than females (z-ratio = 8.95, df = 1, p  < 0.0001; Supplementary Table S6 and Fig. 2B ).

Dogs spending more time alone and participating less frequently in activities and training had higher inattention scores (Supplementary Table S6 and Supplementary Fig. S2B, C ). Dogs that spent alone more than 8 h daily had higher inattention scores than dogs that spent alone less than 3 h (z-ratio = 2.99, df = 1, p  = 0.0097), 3–6 h (z-ratio = 3.85, df = 1, p  = 0.0005) or 6–8 h (z-ratio = 2.49, df = 1, p  = 0.0327) per day. As we hypothesised a priori, there was a difference between dogs that participate in activities and training never/seldom or at least weekly, since dogs participating in activities and training never/seldom had higher inattention scores (z-ratio = 9.52, df = 1, p  < 0.0001). Dogs participating in activities sometimes had higher inattention scores than dogs that trained weekly (z-ratio = 9.28, df = 1, p  = 0.0005).

The dogs with compulsive, aggressive or fearful behaviour also had higher inattention scores (Supplementary Table S6 and Fig. 2C , D, E). Dogs showing high levels of compulsive behaviour had higher inattention scores when compared with dogs showing low (z-ratio = 17.04, df = 1, p  = 0.0005) or moderate (z-ratio = 8.63, df = 1, p  = 0.0005) levels of compulsive behaviour. Dogs showing moderate levels of compulsive behaviour also had higher inattention scores than dogs showing low levels of compulsive behaviour (z-ratio = 12.17, df = 1, p  = 0.0005). Similarly, dogs showing high levels of aggressiveness had higher inattention scores when compared with dogs showing low (z-ratio = 7.50, df = 1, p  = 0.0005) or moderate (z-ratio = 2.72, df = 1, p  = 0.0199) levels of aggressiveness. Dogs showing moderate levels of aggressiveness had higher inattention scores than dogs showing low levels of aggressiveness (z-ratio = 5.93, df = 1, p  = 0.0005). Additionally, dogs showing high levels of fearfulness had higher inattention scores when compared with dogs showing low (z-ratio = 14.86, df = 1, p  = 0.0005) or moderate (z-ratio = 7.90, df = 1, p  = 0.0005) levels of fearfulness. Dogs showing moderate levels of fearfulness had higher hyperactivity/impulsivity scores than dogs showing low levels of fearfulness (z-ratio = 6.85, df = 1, p  = 0.0005).

We have performed the most extensive survey-based study on canine hyperactivity/impulsivity and inattention with over 11,000 dogs, identifying many associated demographic, environmental and behavioural factors. This study is based on the same data as our previous exploration [ 20 ] but we used a more comprehensive and precisive approach here, including complex multivariate models and hyperactivity/impulsivity and inattention scores as continuous variables. We confirmed the behavioural associations observed in our previous study [ 20 ]. In addition, we report novel demographic and environmental associations, observed notable breed differences in the presentation of the phenotypes, and note a significant overlap with the reported risk factors and comorbidities in human ADHD.

Multiple demographic factors were associated with hyperactivity/impulsivity and inattention in dogs, including age, sex and body size. Hyperactivity/impulsivity and inattention were most common in young dogs. Both traits are attenuated with age, but hyperactivity/impulsivity slightly more than inattention. Earlier research has also demonstrated that hyperactive and impulsive [ 22 , 46 , 47 , 48 ] and inattentive [ 21 , 22 , 47 ] behaviour are much more prevalent in young dogs. However, Vas et al. [ 21 ] did not find a significant association between impulsivity and the dog’s age. In our study, hyperactivity/impulsivity and inattention were more prevalent in male than female dogs. This result contradicts earlier studies that have found no significant difference between the sexes [ 48 , 50 ] or have identified females as more hyperactive/impulsive [ 22 ]. Only Vas et al. [ 21 ] found males to be more impulsive than females, but the difference was not statistically significant. Human ADHD is similarly a childhood-onset disease and more common in boys than girls [ 1 , 51 ]. However, the reason for this sex difference is unclear. Girls are more often affected by the predominantly inattentive subtype of ADHD [ 52 ] and show less visible signs of ADHD [ 53 ]. However, girls may require a higher burden of genetic risk factors to manifest ADHD [ 54 ]. A role of steroid hormones has also been proposed, as exposure to high levels of testosterone during gestation might affect the dopaminergic system, and thus potentially predispose boys to ADHD [ 55 , 56 ]. These prenatal hormonal effects have not yet been studied in dogs.

Previous research suggests that small dogs are more impulsive [ 48 ]. In contrast, we observed higher hyperactivity/impulsivity scores in medium-sized and large dogs than in small dogs. Wright, Mills & Pollux [ 48 ] suggested their findings to result from multiple active small terrier breeds. Similarly, our study included several medium-sized working dog breeds, potentially explaining the observed association between higher impulsivity and medium body size. These breeds have been bred for increased activity, alertness and vigilance to maximise their properties as working dogs [ 28 ]. When adjusting the model for body size, the most hyperactive/impulsive breeds in our study included dogs from all sizes and many breeds differed significantly from each other. Thus, the differences between breeds are not explained only by their size differences, and this result also indicates a genetic origin for hyperactivity/impulsivity.

We analyzed hyperactivity/impulsivity and inattention in more than 20 different breeds and found considerable differences. Cairn Terrier, Jack Russell Terrier, German Shepherd Dog, Staffordshire Bull Terrier and Smooth Collie had the highest hyperactivity/impulsivity scores whereas Chihuahua, Rough Collie, Chinese Crested Dog, Miniature Schnauzer and Poodle had the lowest scores. In inattention, Cairn Terrier, Golden Retriever, Finnish Lapponian Dog, mixed breed and Wheaten Terrier had the highest scores, and Border Collie, Poodle, Spanish Water Dog, Shetland Sheepdog and Labrador Retriever lowest scores. Different behaviour traits are valued in breeds used for various purposes, and thus, selective breeding in dogs has influenced their breed-typical behaviour [ 57 , 58 ]. For example, in some working dog breeds, such as German Shepherd Dog and Border Collie, high activity, impulsivity and attention are favoured. These dogs usually have better trainability and working ability due to higher attention spans and reactivity [ 59 ]. On the contrary, these traits are not favoured in breeds which are now preferred as a pet or show dogs, such as Chihuahua, Rough Collie and Poodle, since less active and impulsive dogs are more easy companions in a less active way of life. But, as a side-effect, inattentive behaviour can be enriched in these breeds.

Interestingly, in our study, Smooth Collie was one of the breeds with the highest hyperactivity/impulsivity scores, whereas Rough Collie was one with the lowest scores. Except for coat length, these two breeds are almost identical. However, the use of these breeds nowadays differs. After Lassie movies, the popularity of Rough Collies as companion dogs increased whereas Smooth Collies have never achieved a high level of popularity and are still often used as working dogs and in dog sports [ 60 ]. This might also explain the observed difference in hyperactivity/impulsivity between these closely related breeds. Unfortunately, a comprehensive comparison of our results to earlier findings is difficult as breed groups instead of individual breeds are used in many previous studies [ 22 , 47 , 50 ].

We identified several environmental factors associated with hyperactivity/impulsivity and inattention. Low daily exercise and rare participation in activities and training were associated with higher hyperactivity/impulsivity scores and higher inattention scores, respectively. Ley et al. [ 47 ] found that dogs spending less time inside were more active than dogs spending more time indoors. However, they identified a significant positive correlation between dogs’ age and the time spent inside, suggesting that the observed association might be affected by the fact that older dogs usually spent more time indoors than young dogs. Several studies have shown that dogs that are trained often are less inattentive than dogs trained less frequently [ 21 , 22 , 49 ]. Exercise and enrichment can be ways to fulfil the species-specific needs of dogs. Thus, dogs exercising more and participating more frequently in activities and training can release their energy and frustration in a controlled manner. Therefore, in some cases, high levels of hyperactivity/impulsivity and inattention may be due to limited possibilities to release energy and reduce activity levels. However, it is also possible that owners are not willing to train and participate in activities with inattentive dogs, as training with them can be uncomfortable and unsatisfactory due to the dog’s concentration difficulties. Studies investigating the effects of exercise on human ADHD are sparse and have small sample sizes, but in children with ADHD, a few meta-analyses demonstrate the physical exercise to alleviate hyperactivity, impulsivity and inattention to some extent [ 61 , 62 ].

We observed a novel finding indicating that dogs spending more time alone daily had higher hyperactivity/impulsivity and inattention scores than dogs that spent less time alone. As dogs are social animals, they can be stressed or frustrated when left alone for a long time. This stress and frustration may erupt as hyperactive, impulsive and inattentive behaviour. Dogs generally remain calm and rest during the period of separation from their owners. More extended separation may result in more energetic behaviour and greater physical activity when the owner returns, potentially reflecting dogs’ increased social isolation during the prolonged separation period. Rehn & Keeling [ 63 ] found that after longer separations dogs tended to offer more intense greeting behaviours, with a higher frequency of physical activity and attention behaviour, confirming the effect of time left alone. However, it is also possible that dogs spending more time alone also otherwise get less attention and exercise from their owners. No further conclusions about the relationship between the time spent alone and more hyperactive/impulsive and inattentive behaviour can be drawn, and the causality can only be speculated.

As a novel finding, we discovered that hyperactivity/impulsivity is more common in dogs that are not their owners’ first dogs. As no previous results report a similar association, we can only speculate about the possible relationship between these factors. People may try to choose easy individuals from less active breeds, like companion dog breeds as their first dogs. When they gain experience handling a dog, they may feel more comfortable choosing individuals from more active and challenging breeds, such as herding breeds, as their following dogs. Furthermore, with their first dogs, people may try dog sports and hobbies and if they get excited about the particular sport, they may choose their following dogs from more active and athletic breeds to be more successful in that sport. It is also possible that owners with more than just one dog are more experienced and can better recognise different behaviours, such as hyperactivity/impulsivity and inattention, in their dogs.

Interestingly, both hyperactivity/impulsivity and inattention scores were significantly higher in dogs with high levels of compulsive behaviour, aggressiveness and fearfulness. Impulsivity has been considered in relation to aggressiveness for a long time in dogs [ 13 , 14 , 15 , 19 ]. One recent study discovered a connection between compulsive behaviour and hyperactivity, and between fearfulness and hyperactivity [ 19 ]. Furthermore, Wright et al. [ 48 ] described that impulsivity was more common in dogs reported to have other behavioural problems. Still, they did not declare what these behavioural problems were. However, inattention has been little studied in dogs and these observed comorbidities (between inattention and compulsive behaviour, aggressiveness and fearfulness) have been previously described in dogs only in our previous exploration [ 20 ] of this same but expanded dataset. In this present study, these comorbidities persisted after the inclusion of several demographic and environmental variables in the same multivariate model.

Paralleling our results, ADHD is known to have several behavioural comorbidities, such as autism spectrum disorder, learning impairments, and anxiety and mood disorders in humans [ 1 , 4 , 64 ]. Impulsivity, a key component of ADHD, is also associated with aggressive behaviour [ 8 ]. This can often be classified as impulsive or reactive aggression [ 65 , 66 , 67 ]. Furthermore, obsessive-compulsive disorder (OCD) often co-occurs with ADHD in humans, with both conditions characterised by impaired inhibitory control and deficit in executive function [ 64 , 68 , 69 , 70 , 71 , 72 ].

The comorbid association between hyperactivity/impulsivity and aggressiveness, fearfulness and compulsive behaviour may refer to shared underlying neurobiological pathways and brain structures involved in these traits. ADHD and impulsive behaviour are associated with deficits in the frontostriatal circuit and abnormal levels of activation in, for example, prefrontal cortex (PCF), anterior cingulate cortex (ACC) and striatum [ 64 , 70 , 73 ]. Similarly, OCD is also characterised by abnormal frontostriatal circuit activity and likewise, involves the striatum, PFC and ACC [ 71 , 72 , 74 , 75 ] Besides ADHD and OCD, aggressiveness also involves the brain reward system and neurological pathways involved in aggressiveness similarly connect to PFC and striatum [ 65 , 67 ]. Finally, fear and anxiety are also associated with activity in PFC and ACC [ 76 , 77 ].

Our results indicate that different breeds may be useful in modelling the different presentations of ADHD. Cairn Terrier could be a suitable model for ADHD as it has a high mean score in both hyperactivity/impulsivity and inattention traits, together with compulsive behaviour and aggression comorbidities. In contrast, Labrador Retriever had a low mean score in both traits and they also display comorbid behaviours fearfulness, aggression and compulsive behaviour very rarely. Spanish Water Dog had a low mean score in inattention, but a high mean score in hyperactivity/impulsivity, together with comorbidities, such as fearfulness. In contrast, Chinese Crested Dog had a high mean score in inattention, but a low mean score in hyperactivity/impulsivity as well as high levels of fearfulness, aggression and compulsive behaviour. Finally, within-breed studies would be helpful in revealing the genetic and biological factors associated with hyperactivity/impulsivity and inattention.

This study has limitations. Our analysis is based on a questionnaire and owners’ participation in the study was voluntary. Questionnaires can be an effective way to collect data as their reliability has been useful in behavioural science and questionnaire answers are strongly linked to the behaviour of the animals [ 34 , 37 ]. However, questionnaires can be subjective. Our data is a self-selected convenience sample and may not represent the overall dog population in Finland. Due to missing data, several dogs were excluded from the analyses and thus, future studies should aim to collect more complete data. Finally, some of the breeds we studied included both working and show lines and we could not separate the lines within the breeds. In our future studies, we aim to collect information about the line of the dog to assess the possible behaviour differences between the show and working lines.

In conclusion, we show that canine hyperactivity/impulsivity and inattention are associated with several demographic, environmental and behavioural factors. Our results also suggest that these traits have a strong genetic basis. Furthermore, our results reinforce the dog as an appropriate, up-and-coming animal model of ADHD. Hyperactivity/impulsivity and inattention were more common in young and male dogs, and the same age and sex effects are well established in human ADHD as well. Additionally, similar behavioural comorbidities in canine hyperactivity/impulsivity and inattention and human ADHD strengthen the hypothesis of the shared neurobiological pathways behind these traits in both species. Furthermore, the similarities in genetics, physiology and living environment between dogs and humans make the dog a more intrinsic model to ADHD than, for example, rodents. Therefore, understanding the factors that affect canine hyperactivity/impulsivity and inattention can benefit not only recognition and management of these traits in dogs but also human ADHD research.

Data availability

The anonymized data is available as Supplementary material in the article by Salonen et al. [ 20 ].

Code availability

The R environment is freely available at https://www.r-project.org/ . Additionally, all the R packages utilised for the analyses are freely available at https://cran.r-project.org/web/packages/available_packages_by_name.html . Specific R scripts used for the analyses are available from the corresponding author on reasonable request.

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Acknowledgements

We thank all dog owners who participated in this study with their dogs. We acknowledge Katriina Tiira for her contribution to the canine behavioural questionnaire development, and Joona Lehtomäki for his assistance in determining the urban environment score variable. This study was partially funded by the Academy of Finland (308887), the ERCStG (260997), ERA-NET NEURON, the Jane and Aatos Erkko Foundation, Doctoral Programme on Veterinary Medicine. H.L. is a member of HiLIFE. M.S., S.M., E.H. & H.L. are members of the Helsinki One Health.

Author information

These authors contributed equally: Sini Sulkama, Jenni Puurunen.

Authors and Affiliations

Department of Veterinary Biosciences, University of Helsinki, Helsinki, Finland

Sini Sulkama, Jenni Puurunen, Milla Salonen, Salla Mikkola, Emma Hakanen, César Araujo & Hannes Lohi

Department of Medical and Clinical Genetics, University of Helsinki, Helsinki, Finland

Folkhälsan Research Center, Helsinki, Finland

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Contributions

Study design: M.S., S.S., H.L., J.P., E.H. and S.M.; data collection: M.S., S.S., J.P. and C.A.; resources: H.L.; data analysis: M.S. and S.S.; figure preparation: M.S.; funding acquisition: H.L. and S.S.; manuscript writing: S.S. and J.P., manuscript editing and review: H.L. S.M., E.H. and M.S.; Supervision: H.L.

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Correspondence to Hannes Lohi .

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Competing interests.

The authors declare no competing interests.

The data were collected according to the Finnish legislation ( https://www.finlex.fi/fi/laki/ajantasa/1999/19990523 ) before the onset of the GDPR. A specific ethical approval was not needed at that time for academic research studies as this study focused on dogs and not humans or dog owners. Only names and addresses of dog owners were collected for contacting them in subsequent studies and for calculating the urban environment score. We informed the dog owners that participated in the study is entirely voluntary and confidential. We emphasised that the data will be used only for scientific purposes. We received informed consent from all participants.

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Sulkama, S., Puurunen, J., Salonen, M. et al. Canine hyperactivity, impulsivity, and inattention share similar demographic risk factors and behavioural comorbidities with human ADHD. Transl Psychiatry 11 , 501 (2021). https://doi.org/10.1038/s41398-021-01626-x

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Received : 03 March 2021

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Published : 01 October 2021

DOI : https://doi.org/10.1038/s41398-021-01626-x

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USM Graduate Student Writes First-Author Publication for Bee Research

Thu, 05/16/2024 - 01:05pm | By: Van Arnold

University of Southern Mississippi (USM) graduate student Kristin Robinson’s research on how bees are affected by climate change has been published this week in a special issue on Insect Conservation Behavior in the journal Frontiers in Ecology and Evolution.

Robinson is a doctoral candidate in USM’s School of Biological, Environmental and Earth Sciences (BEES). Her paper, titled “Stingless bee foragers experience more thermally stressful microclimates and have wider thermal tolerance breadths than other worker subcastes” addresses how climate change impacts key ecosystem services that insects provide, such as pollination. Robinson works in the lab of Dr. Kaitlin Baudier, Assistant Professor in the School of BEES.

“It is fairly straightforward to estimate what temperatures cause bees distress, but in order to accurately put performance data together with future temperature projections, we need to understand the behavioral strategies that pollinators use in their environment. In her study, Kristin did just this,” said Baudier.

A native of Minneapolis, Minn., Robinson immersed herself in biology research while pursuing an undergraduate degree at the University of Minnesota. The fieldwork associated with the species highlighted in her research has been conducted in Costa Rica. She acknowledges the honor of being published as a first author on a paper for the first time.

“Seeing the payoff of this first publication from my dissertation research is incredibly rewarding after putting in the work to get to where I am now,” said Robinson. “Of course, I am also very grateful for all the people who have supported me personally and professionally over the years to pursue my education and research.”

She added, “As a graduate student, I think we are all often plagued with insecurities about how much we still have to learn and the feeling that we are unfinished scientists in the making. While that is very much true because we are still students after all, being able to publish my work does help me to see my research from an outside perspective and realize that I am doing work that is valuable and appreciated.”

Last year Robinson received a Charles Michener Bee Research Grant from the North American Section of the International Union for the Study of Social Insects to support her follow-up work investigating how division of labor interplays with the desiccation limitations of the same species of stingless bee.

Baudier stressed the importance of Robinson’s work which helps scientists understand how a tropical pollinator may fare in the future as the temperature rises. The research also sheds new light on what factors shape the evolution of within-colony thermal tolerance variation.

“Kristin is committed to pursuing answers to important questions related to insect conservation. She is a knowledgeable field biologist and an all-around bright early career researcher,” said Baudier. 

Categories: Research

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