In Are We Smart Enough to Know How Smart Animals Are? Frans de Waal presents a fascinating history of the study of animal behavior and cognition. De Waal, who says his love of animals dates to his childhood, is a worldrenowned primatologist and ethologist and director of the Living Links Center at the Yerkes National Primate Research Center. We asked him to shift gears and give us his take on the canine mind.

Bark: Konrad Lorenz (co-founder of your field) wrote Man Meets Dog in 1954. And while it is still one of the best, if slightly flawed, books on canine behavior, why did it take so long for ethologists, and other researchers, to to study dog behavior?

Frans de Waal: Dogs were (and are) considered imperfect subjects of study because they are “unnatural.” Many ethologists, including Lorenz, feel that natural behavior under naturalistic conditions is what we should focus on, and the dog is a product of artificial breeding. Lorenz liked all animals, however, and so couldn’t resist describing his dog stories, and we should all be grateful.

Clearly, the dog is a mammal with many typical mammalian tendencies, so now scientists are finally seeing that the fact that they are domesticated also has advantages. For example, they are eager to work with us, they are generally not dangerous, they are smart, they have empathy. Lots of great things can be done with them. And they are easier to work with than other large mammals, such as apes and dolphins.

Bk: Can you give an example of how other species, including dogs, demonstrate empathy?

FdW: American psychologist Carolyn Zahn-Waxler sought to determine at what age children begin to comfort family members who sobbed or cried “ouch.” It turns out that children do so at one year of age. In the same study, Zahn-Waxler accidentally discovered that household dogs react similarly. Appearing as upset as the children by the distress-faking family members, the dogs hovered over them, putting their heads in their laps with what looked like great concern. This work has recently been repeated in different studies, more focused on the dogs themselves, and it is clear that these animals show empathic concern for humans.

The ancestor of the dog, the wolf, probably behaves the same. If “man is wolf to man,” as Thomas Hobbes liked to say, we should take this in the best possible way, including a tendency to comfort the whimpering and help the needy. This insight, of course, would undermine much of political philosophy based on Hobbes’ dog-eat-dog view of nature.

Bk: Do you think human bias has played a part in some of the canine cognitive studies?

FdW: At first, dogs were rated as more intelligent than even apes and wolves because they followed the direction of human pointing (at a bucket with food), whereas apes and wolves ignored human directions. Then it was found that wolves raised in a human home will act more like dogs, following human pointing, suggesting that the earlier failures with wolves were probably due to lack of bonding and attention. The same probably applies to the apes. Now, dogs are seen not just as smart but rather, as finely in tune with the species that bred them.

They have a special bond with us, as also reflected in the oxytocin studies, which show that human-dog contact increases this “cuddle” hormone in both. The dog is perhaps the only animal that performs at its peak when tested by humans, whereas many other animals are not so into us, hence need to be tested in different ways. This is yet more proof that cognitive testing of animals always needs to take into account what kind of animal we are dealing with: we need to find the most species-appropriate way.

Bk: In contrast to behaviorism’s reward/ punishment model, ethology views animals as “seeking, wanting and striving.” Why do you feel the latter is a more productive way to look at animals?

FdW: The behaviorists (followers of B. F. Skinner) totally overlooked natural animal tendencies. Trying to explain all behavior on the basis of reward and punishment, they could not explain why you can train a dog to fetch, but not a rabbit or a goat.

Predators are obsessed with small moving objects, which we see every day in our dogs as well as cats. Their interest sets up a learning situation where they are going to absorb many lessons about how to catch these moving objects, how to trick them, how to outsmart them. Dogs eagerly learn all of those things.

Reward and punishment are only small parts of the story; their natural hunting instinct is, in fact, the driver of the process. This is where behaviorism failed. It had some good ideas, many of them applicable to animal training, but its perspective was far too narrow as it lacked attention to natural tendencies and the evolution of behavior.

Bk: Why do you think Darwin used dogs to illustrate emotional continuity?

FdW: Darwin was a dog lover, and he knew that to get his message across about the continuity between human and animal emotions, the dog would be the easiest way to communicate. Darwin mostly worked on the expression of emotions (it’s hard to know what animals feel, but we can at least document how they signal various states, such as fear, submission, anger, affection). Of course, the dog is very expressive with its postures, facial expressions, tail-wagging, growling and so on. Darwin knew that most people could relate to all of this, and would have more trouble if he described other species that people have less exposure to.

Bk: In terms of an evolutionary advantage, how important is it for a species to have self-awareness, or theory of mind?

FdW: These capacities require large brains. In terms of recognizing oneself in the mirror or understanding what others know, the champion species are apes, dolphins, elephants and perhaps also the corvids (crow family). This doesn’t mean that dogs lack them. They probably have similar understanding, but not as fully expressed.

The more complex the societies of a species, the more demands there are on cognition, and perhaps canines do not need social understanding at the level of an ape or dolphin. I feel we need to judge animals on what they are good at and what they need to know to survive. In this regard, canines have lots of specialized skills, often related to their sense of smell, their pursuit of prey, their need for tight cooperation and so on. This is where we should test them out, and probably find remarkable skills.

Bk: Clearly, emotions are important to the understanding of behavior; how do they relate to and inform one another?

FdW: In my book, I left emotions out on purpose because I felt it would muddle things. But there can be no studies of cognition without attention to the emotions, and vice versa. The two go hand in hand. In our famous capuchin monkey experiment with the grape and the cucumber, for example, you can see not only that the monkeys judge what they get relative to what others get, but also their strong emotional response. You cannot study the one and ignore the other.

Humans use both words and the intonation of speech to decipher the meaning of language, and it turns out that our dogs do, too. In a research paper called “Neural mechanisms for lexical processing in dogs” scientists investigated how dogs process the meaning of language. They found that dogs’ brains have even more in common with humans’ brains than previously thought. (It’s not clear when we will collectively stop being surprised by this, but I hope we always remain excited about new evidence to explain why we feel that dogs are kindred spirits.)

In this study, dogs who have been trained to remain still while their brain activity is recorded listened to recordings of their trainers talking. There were four types of recordings: 1) words of praise spoken with intonation typically associated with praise, 2) words of praise spoken with a neutral intonation, 3) neutral words spoken with intonation typically associated with praise, and 4) neutral words spoken with a neutral intonation.

Researchers analyzed the brain activity of the dogs in response to each of the recordings, and came to several conclusions about the way that dogs respond to words and the intonation of human speech. The dogs processed the vocabulary in the left hemisphere of their brains, which is where humans also process the meaning of words. The dogs processed the intonation of the words separately, in a different region of the brain. Just as humans do, dogs processed the intonation of human speech in the right hemisphere of their brain. Dogs also process sounds that convey emotion without words in this same region of the brain’s right hemisphere.

Dogs process both words and the intonation of human speech to decipher meaning. Just as humans do, they process these two aspects of speech separately, then integrate them to determine the full meaning of what was said. Only the praise that was spoken like praise—higher pitched than normal speech and with more variation in pitch—activated the reward centers of dogs’ brains. Though they may understand words of praise said in any manner, it only makes dogs happy to hear us praise them when we do it with proper feeling.

This research does more than reveal yet another similarity in the way that human and dog brains process information. It also suggests that the ability to connect a word to a meaning did not develop with the evolution of spoken language. Rather, it is a more ancient ability that can be made use of in the context of the human-dog relationship to link specific sounds to specific meanings.

The take away messages from this research are that dogs process two parts of spoken language—words and intonation—the same way that humans do and if you want to make your dogs happy, you have to praise them like you mean it!

Not surprisingly, a study published July 29, 2016 found that the English Bulldog no longer retains enough genetic diversity to correct life-threatening physical and genomic abnormalities. This means breeders cannot use the established population of purebred dogs to reverse the trend in extreme and painful exaggerations such as crippling dwarfism and respiratory deformities - traits that uninformed pet-owners find appealing.

In the early 1800s Bulldogs were trained for bull-baiting, a particularly cruel and vicious sport. In 1835 the Royal Society for the Prevention of Cruelty to Animals convinced Parliament to enact the first animal cruelty law for the protection of domestic animals, including outlawing bull baiting. 

As such, the Bulldog had outlived its usefulness. Like the pre-19th century Wolfhound that disappeared with the eradication of wolves in the British Isles, and the Tumbler whose demise was the invention of hunting firearms, the Bulldog was destined for extinction.

English Bulldog from 1890

But it was not to be. Beginning about 1840, the Victorian dog fancy's unabashed sentimentality was a catalyst for saving even the most formidable working breeds from their inevitable demise. Like many others, such as the Dachshund and Mastiff, Bulldogs went from working hard to hardly working.

Utility dogs were "refined" and transformed to fill jobs they weren't originally bred for - as show dogs and companions. Altered physical and behavior characteristics along with decreased levels of aggression were more compatible for their augmented duties as house pets.

English Bulldogs from 1920s

Beginning in the late 1890s, Bulldog breeders (and other breeders as well) selected small groups of genes from a diverse genome and created new breed-types. They were in effect increasing the odds that genetic anomalies would more likely be expressed to bring out exaggerated traits, like the Bulldog's baby-like face, corkscrew tail and affable personality.

As "desirable" aesthetic traits were selected for, other genetic variants including beneficial genes that contribute to overall health were eliminated from the gene pool, never to be reclaimed.

In the last few decades the most exaggerated traits in the Bulldog - the extreme brachycephalic skull and deformed skeleton- have become increasingly pronounced because naive consumers want that type of dog and consequently that's what many breeders select for.

Driven by economics, fashion, and uninformed decisions, breeders and buyers either ignore or are unaware of the genetic problems that have spread throughout the population.

The demise of the breed may not be a good thing for Bulldog-lovers, but it will thankfully put an end to the malformed and painfully crippled modern Bulldog we recognize today.

The good news is that some breeders are intent on bringing back the "Olde-Fashioned-Bulldogge". 

The common wisdom that dogs can smell fear doesn’t give dogs full credit to the nuances of their ability to sense emotion through their noses. A recent study titled “The dog nose “KNOWS” fear: Asymmetric nostril use during sniffing at canine and human emotional stimuli” examined dogs’ tendencies to sniff various substances with the right or the left nostril. Exploring this side bias may seem like looking at random details, but the side of the nose used to sniff something tells us a lot about the dog’s emotional reaction to the odor. The use of one side of the body indicates a differential use of one side of the brain or the other, which is a clue to the dog’s emotions.

The left side of the brain processes more positive emotions such as happiness and excitement as well as stimuli that are familiar. The right side of the brain tends to take over when a dog is processing negative emotions such as sadness or fear as well as novel stimuli. In general, the right side of the body is controlled by the left hemisphere of the brain and vice versa. However, the nose is an exception; the right nostril sends information to the right side of the brain to be processed and the left nostril sends its information to the left side. The findings of this study suggest that the pathways used to process various olfactory stimuli are dependent on more than just whether they elicit negative or positive feelings.

Eight odors were tested—four from dogs and four from humans. The four human odors were collected as sweat from donors who were joyful, fearful, physically stressed, or in a neutral situation. The joyful and fearful states were elicited by movies, and the physical stress odor was collected after donors ran for 15-minutes. The four canine odors were collected from dogs who were happy following a play session with the guardian, stressed by isolation in an unfamiliar place, disturbed by a stranger approaching the car, and dogs who were asleep. The dogs who “donated” odors were different from the dogs whose sniffing behavior was studied.

To further explore the phenomenon of side bias in sniffing, the guardians of the dogs in the study filled out a questionnaire related to each dog’s temperament. During the study, dogs were led to a video camera under which was mounted a Q-tip saturated with various odors. The videos captured the dog’s sniffing behavior so that it was possible to determine a laterality index for each dog for every odor based on the amount of time spent sniffing with each nostril. A laterality index of 1.0 indicated exclusive use of the left nostril and negative 1.0 indicated exclusive use of the right nostril. Dogs’ cardiac activity was also recorded during the tests of each odor.

I’m sure it’s the science geek in me, but I got a kick out of reading the sentence, “Results for nostril use are shown in Figure 2.” Three of the odors elicited consistent sidedness in nostril use and five of them did not. Dogs more frequently used the right nostril to sniff the canine isolation odor. They more frequently used the left nostril to sniff the human fear odor and the odor from human physical stress.

There were two ways in which the results of the questionnaire were correlated with the laterality pattern for a particular odor. The higher a guardian ranked the dog’s fear/aggressiveness to other dogs, the more likely that dog was to use the right nostril for sniffing the disturbed canine odor. This suggests that individual differences in emotional arousal and perhaps even in temperament influence asymmetries in sniffing behavior. Dogs with higher scores for predatory behavior used the left nostril more for sniffing the odor that came from physically stressed humans. This makes sense when we consider that it is structures in the left side of dogs’ brains that are involved in predatory behavior.

Dogs’ brains are every bit as amazing as their noses, as research about both of them reveal!