Bonding and Stress

Bonding in animal behavior is a biological process in which individuals of the same or different species develop a connection. The function of bonding is to promote cooperation.

Parents and offspring develop strong bonds—the former takes care of the latter, and the latter trusts the teachings of the former. As a result of filial bonding, offspring and parents or foster parents develop an attachment. This attachment ceases to be necessary once the juvenile reaches adulthood, but may have long-term effects on subsequent social behavior. Among domestic dogs, for example, there is a sensitive period from the third to the tenth week of age, during which common contacts develop. If a puppy grows up in isolation beyond about fourteen weeks of age, it will not develop the relationships considered normal for species and population.

Males and females of social species develop strong bonds during courtship motivating them to care for their progeny, so they increase the probability of the survival of 50% of their genes.

Social animals develop bonds by living together and having to fend for survival day after day. Grooming, playing, reciprocal feeding, all have a relevant role in bonding. Intense experiences do too. Between adults, surviving moments of danger together is strongly bonding.


The strongest bonds originate under times of intense experiences (photo by Cuttestpaw).

Behavior like grooming and feeding seems to release neurotransmitters (e.g., oxytocin), lowering the innate defensiveness and increasing the odds of bonding.

We often mention bonding together with imprinting. Even though imprinting is bonding, not all bonding is imprinting. Imprinting describes any phase-sensitive learning (learning occurring at a particular age or a particular life stage) that is rapid and apparently independent of the consequences of behavior. Some animals appear to be preprogrammed to learn about certain aspects of the environment during particular sensitive phases of their development. This learning is preprogrammed, in the sense that it will occur without any visible reinforcement or punishment.

Our dogs, in the domestic environment we offer them, develop bonds in various ways. Grooming, resting together, collective barking, and playing and chasing intruders are efficient at creating bonds. Their bonding behavior is by no means restricted to individuals of their species. They bond with the family cat as well and with us, humans.

Bonding is a natural process that will inevitably happen when individuals share responsibilities. Looking into one another’s eyes is only bonding for a while, but surviving together may be bonding for life—and this applies to all social animals, dogs and humans included.

We develop stronger bonds with our dogs by solving problems together rather than by just sitting and petting them. These days, we are so afraid of anything remotely connected to stress that we forget that the strongest bonds originate under times of intense experiences. A little stress doesn’t harm anyone, quite the contrary. I see it every time I train canine scent detection. The easier it is, the quickest it will be forgotten. A tough nut to crack, on the other hand, is an everlasting memory binding the parties to one another.

One of the most exciting scientific discoveries of the latest is on epigenetics. Epigenetics is the study of heritable changes in gene activity not caused in the DNA.

Stress hormones seem to boost an epigenetic process either increasing or decreasing the expression of particular genes. Stress hormones change specific cells of the brain that help memories to be easier retained.

We need to be careful. The term stress is dangerously ambiguous. “Stress is a word that is as useful as a Visa card and as satisfying as a Coke. It’s non-committal and also non-committable,” as Richard Shweder says.* I’m talking about stress in a biological sense, the response of the sympathetic nervous system to some events, its attempts at re-establishing the lost homeostasis provoked by some intense event.

Being an evolutionary biologist, when contemplating a mechanism, I always ask: “What is the function of that? What is it good for?” A behavior can originate by chance (most do), but if it does not confer the individual some extra benefits as to survival and reproduction, it will (most likely) not spread into the population.

Asking the right question is the first step to getting the right answer. Never fear to ask and reformulating your questions. At one point, you’ll have asked the question that will lead you to the right answer.

Why do unpleasant memories seem to stay with us longer than pleasant ones,sometimes even for the rest of our lives? 

Situations of exceeding anxiety and stressful, intense experiences create unpleasant memories. It is important, if not crucial, to remember situations that might have hurt us seriously. It makes sense that the stress hormones should facilitate our retaining the memory of events occurring under stress.

Stress hormones do bind to the particular receptors in the brain that enhance the control of the epigenetic mechanisms involved in remembering and, hence, in learning. They do boost the epigenetic mechanisms that regulate the expression of the genes crucial for memory and learning.

Not all stress boosts learning. Too much stress produces the opposite effect. There is a difference between being stressed and stressed out. When we experience far too much stress, our organism goes into an alarm mode where survival has the first and sole priority, and memory formation decreases. Chronic stress does not promote learning either.

Bottom line: we need to be nuanced about stress. Events causing healthy stress responses are necessary for enhancing attention to details, the formation of memory, the creation of bonds, and learning—and too much stress works against it.


* Burnout vs Depression – Human Stress

Featured image: Dog and Child in a bonding moment (photo by Elena Shumilova).

Superstitious Behavior in Animal Training


Superstitious behavior is behavior we erroneously associate with particular results. Animals create superstitions as we do. If by accident a particular stimulus and consequence occur a number of times temporarily close to one another, we tend to believe that the former caused the latter. Both reinforcing and inhibiting consequences may create superstitious behavior. In the first case, we do something because we believe it will increase the odds of achieving the desired result (we do it for good luck). In the second case, we do not do something because we do not want something else to happen (it gives bad luck).

In 1948, B.F. Skinner recorded the superstitious behavior of pigeons making turns in their cages and swinging their heads in a pendulum motion. The pigeons displayed these behaviors attempting to get the food dispensers to release food. They believed their actions were connected with the release of food, which was not true because the dispensers were automatically programmed to dispense food at set intervals.

Timberlake and Lucas concluded in 1985 “[…] that superstitious behavior under periodic delivery of food probably develops from components of species-typical patterns of appetitive behavior related to feeding. These patterns are elicited by a combination of frequent food presentations and the supporting stimuli present in the environment.”

We should be very careful when reinforcing any desirable behavior the animal we train shows us. If the reinforcement happens to coincide with other more or less accidental stimuli, we may be creating superstitious behavior. We may create superstitious behavior with any reinforcement, but probably food is the most liable to do it.

The same goes for inhibitors.1 We should always bear in mind that we never reinforce or inhibit an individual but rather a behavior.2


This can be superstitious behavior: the dog believes that if it barks long enough at the door, someone will open it because it has happened before. Many CHAP (Canine Home Alone Problems) are not even remotely connected with anxiety as many dog owners erroneously presume.

In one of our Guinea Pig camps, we saw some Guinea pigs displaying superstitious behavior. One of them would place a paw on the tin containing the target scent and would swing its head repeatedly in the direction of the trainer. The piggy created this superstition because the trainer presented the reinforcers (“dygtig” and food) when it swung its head and not when it placed the paw on the tin right after sniffing the target scent. It did not take many repetitions before the animal had created an erroneous association. Another piggy would walk over the tin if it didn’t get a reinforcer right away.

Superstitious behavior is extremely resistant to extinction. Skinner found out that some pigeons would display the same behavior up to 10,000 times without reinforcement. Displaying a behavior expecting a reinforcer, and receiving none, increases persistence. It’s like we (as well as other animals) feel that if we continue long enough the reinforcement will happen sooner or later.

As always, being an evolutionary biologist, the first question that comes to my mind is, “what conditions would favor the propagation of superstitious behavior?” Making correct associations between events confers a substantial advantage in the struggle for survival. That is what understanding (or adapting to) one’s environment means. The benefits of getting one association right outweigh the costs of making several wrong associations so much that natural selection favors those who tend to make associations rather than those who do not—and that’s why superstitious behavior is highly resilient to extinction.
1“Inhibitors” were earlier called “punishers.” I coined the new term in “The 20 Principles All Animal Trainers Must Know.
2 In Abrantes 2013 and 2016.

Abrantes, R. (2013) The 20 Principles That All Animal Trainers Must Know. Wakan Tanka Publishers.
Abrantes, R. (2016) Animal Training My Way, Wakan Tanka Publishers.
Skinner, B. F. (1948). Superstition in the pigeon. Journal of Experimental Psychology, 38, 168-172.
Timberlake, W and Lucas G. A. (1985). The basis of superstitious behavior: chance contingency, stimulus substitution, or appetitive behavior? J Exp Anal Behav. 1985 Nov; 44(3): 279–299.

Featured image: Superstitious behavior is easy to create and extremely difficult to extinguish (photo from

The Misadventures of Bongo


In 1994, I created Bongo to illustrate the various situations dog owners and dogs get into and how to get out of them the best possible way. My objective was to explain and illustrate that many dog problems (maybe most) were the result of misunderstandings between us and them. If we spoke a better “Doguese,” we could certainly avoid the worst troubles. I paired up with Henriette Westh, a brilliant Danish illustrator, and she gave Bongo more than a form; she gave him a character of his own.

Bongo is a nice, friendly and naughty English Cocker Spaniel (orange roan, the original drawings were in color) with his own mind. He’s a good dog and loves his family very much, but he gets often in trouble, mostly because of misunderstandings as you can see in “Bongo Home Alone.”

“Bongo Home Alone” was first published in 1994 in my book “Hunden, ulven ved din side” (Borgen Publishers). Coincidentally, the editor of the book was none other than Henriette’s brother, Poul Henrik Westh. The book never appeared in English, but Bongo did.

Enjoy this bit of history and nostalgia and have a good laugh!



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Featured image: Bongo Home Alone (Illustration by Henriette Westh)

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A Dog’s Self-Respect


Did she cheat me? Did she manipulate me? Was it a proof that my English Cocker Spaniel had a sense of self-respect, that dogs behave intelligently?

It happened long ago, but I still think about it, trying to find a plausible and scientifically correct explanation. My dogs have always been fun dogs, independent and skillful, but manipulative and naughty at the same time. It’s my fault. I’ve brought them up to be that way. I trained them because at the time (the beginning of the 1980s) I was keen on demonstrating that there were other ways of training dogs than the traditional, mostly compulsory and often forceful methods of the old school. Since I believed (and still do) that the best way to have someone change is not by forcing, persuading or convincing, but rather by showing attractive results, I trained my dogs to help me in this quest, and none more than Petrine, my female, red English Cocker Spaniel did so.

At the time, there was a very popular dog training series on TV called “No Bad Dogs the Woodhouse Way” with the unforgettable Barbara Woodhouse. Those of a certain age will chuckle nostalgically when they hear inimitable “walkies.” Mrs. Woodhouse, born in 1910, was a charming, efficient lady who loved animals. She was not mean; it was just her methods that were forceful to say the least. Does this sound familiar? History repeats itself, as we well know!

Instead of attacking her and her methods personally, or trying to argue for ways I thought were better, I found a more convenient strategy: to channel the interest in dog training that Mrs. Woodhouse generated and present my own way as an alternative. Of course, I had to show results. I had to be able to teach the dogs the same Mrs. Woodhouse so efficiently taught them. If I was successful, and my methods were not only as efficient but more attractive, they would win the public’s favor. If I couldn’t achieve the same results as her, my way would not win. I went for it, confident that I could make dogs as “obedient” as Mrs. Woodhouse did, but using my own methods. To allow for an obvious comparison, I even used the terminology of the time, which I later felt entitled to change when my first book came out in 1984: from there on a “command” became a “signal,” “obedience” became “cooperation,” and “praise” became a “reinforcer.”

So, Petrine and I did a lot of “obedience” training, even if we weren’t too keen on the fastidiousness of the process. We trained using motivation, treats, facial expressions as reinforcers, the word “dygtig,” later to be called a semi-conditioned verbal reinforcer, and sometimes a whistle as a conditioned positive reinforcer (the precursor of the clicker); and together we won several obedience competitions.

At the time, you didn’t see many Cockers competing, and our victories did help to prove my point, but our achievements weren’t exactly a big surprise. They were more like appetizers. What really did it was when we won a hunting-dog competition. That caused quite some stir in the dog-training community of that time because we beat all the smart, green clad hunters with their pointers and the like. At the time, it was unthinkable that an English Cocker Spaniel (not only red, but female too!) and a longhaired, bearded, young fellow (in worn-out Levi’s and clogs just to top it off) could beat the establishment. Well, we did! That day of fame and infamy set me on a career path I could never have imagined. Training in a new way, the “psychology rather than power” way rather than the Woodhouse way, we made it into newspapers, magazines, TV and radio, and to be on TV was a big thing at the time. Inevitably, we were heroes for some and villains for others, but my message had been conveyed as the first edition of my first book, entitled (of course) “Psychology Rather Than Power,” which showed a completely different way of training dogs based on ethology and the scientific principles of animal learning, sold out in three months. It was a victory for psychology rather than power in more than one way, as it also proved my point that showing results works better than arguing, persuading, convincing or forcing.

Petrine was indeed an amazing dog. She taught me most of the important things I know about dogs, but she also taught me about life, respect and affection. As I said before, I trained her because it was necessary, but I must confess that I never liked the training as much as the interaction. Training was definitely secondary to having a good relationship. Therefore, I always encouraged and reinforced any behavior that showed initiative, independence, and her resolving problems her own way. This was (and is) my philosophy of education for any species. I think of my job as an educator as like being a travel guide, providing my students with opportunities to develop, to learn how to deal with their environment, to stand out from the crowd and not be just a self-denigrating face, but to make of themselves whatever they choose. If my dogs found ways to circumvent the rules and succeeded (that is what I call good canine argumentation and reasoning), I would reinforce that even at my own cost. In other words: I have always reinforced sound argumentation and conclusions consistent with their premises, even though they might have gone against my wishes and, as the good sportsman, my father educated me to be, when a better opponent on a better day beats me, I accept defeat gracefully. I applied the same philosophy to the education of my son.

When Daniel was little, we traveled a lot together. I always thought traveling, experiencing other ways of thinking and having other stances on life were good antidotes to narrow-mindedness and all that comes with it. On one occasion, we arrived at a guesthouse after a long journey and Daniel, by then about 9 or 10 years old and already an experienced traveler, quickly assessed the situation.

“OK, we have only one little bed,” he said.

“Yes, so I see,” I replied, whilst removing my heavy backpack, trying not to lose the car keys or spill our cokes.

“I have 50% of your genes, and when I have kids, they’ll have 25% of your genes, right?” he asked rhetorically.

“For sure,” I said, amazed at what a kid could learn just by accompanying his daddy to talks and seminars whilst quietly drawing pictures at the back of the room.

“So if you want me to pass 25% of your silly genes to my kids, you have to take good care of me, right?” again a rhetorical question.

“Yes, absolutely,” I answered.

“OK, so I take the bed, and you sleep on the floor,” he concluded.

I slept on the floor.

Petrine, the red, female English Cocker Spaniel was indeed one of a kind. I remember one day I had decided to invite guests for dinner and prepared a roast beef to serve them. It was no mean feat considering my extremely limited culinary skills. I was in the living room surveying the table when I glanced towards the kitchen, and my eyes registered a sight that caused instant paralysis of every muscle in my body, including my jaw, which gaped open as I recollect.

Next to the kitchen table, where I had placed the fruit of my hard labor, the once-in-a-lifetime masterpiece, my roast beef, stood Petrine. That in itself is not reason enough to make me stop breathing and incite a serious and irreversible heart-attack you may think, and you’re right, but add to that Petrine holding my roast beef in her mouth and I think you will begin to understand the cause of my instant, full body paralysis. For a moment that seemed interminable, we stood there looking at one another, me, drop-jawed and paralyzed from head to toe, and Petrine with her deep brown eyes staring at me intensely, roast beef in mouth.

If I was paralyzed, Petrine certainly was not. She began to walk towards me with a swift, self-confident, elegant pace, not once averting her gaze from mine. I merely stared in disbelief at her approach with the roast beef. Without stopping, she trotted around me in a perfectly calculated circle and sat right next to my left leg, lifting her head and the roast beef towards me, her eyes still fixed on mine.

I think I took longer to react than I normally would on this type of occasion, but I managed to bend down, take hold of the dummy (read roast-beef) and give the signal “Tak” (read release). I know I managed it because I remember trying to wipe away Petrine’s teeth marks from the roast beef and placing it on a plate on the table ready to serve to my guests. I also remember that, even though my paralysis had only been momentary, my brain was still not fully functioning, as the next I heard was a barely perceptible whine from Petrine. I looked down to find her gazing up at me, wagging her tail and all lower body as cockers do. She was right, and it was good of her to remind me. I was failing in my duties. “Free,” I said, and, as swiftly, as elegantly and as self-confidently as she had brought the roast beef to me, she went off to perform some other of her daily chores. It had all been just another episode among the many life presents us with. No more, no less— or so it seemed to her.

It was only once the guests had gone, the kitchen tidy and Daniel in bed that, sitting on my porch and enjoying a well-deserved glass of Portuguese “vinho verde,” I cast my mind back to the Petrine episode. What had been going on?

As I told you, my philosophy of education encourages determination and reasoning, and Petrine was good at that. She realized that she had been caught in the act. She had several options: one, to drop the roast beef and show submissive behavior (active and/or passive), which would have been accompanied by a “Phooey” from me, an ugly face and a very assertive tone of voice; two, to scoff as much of the roast beef as she could before I caught her, which wouldn’t have taken long considering I was no more than 6 meters (20 feet) away; three, to run with the roast beef, which she could have done, but I would inevitably have caught up with her. And, of course, she also had the option that she chose, which is not one I would have thought of myself. Why did she choose that option? All things considered, I believe it was the best option open to her, but what went through her head when she chose to do so, I would pay a handsome fee to know for sure.

None of my (attempted) scientific explanations succeeds in convincing me fully. Having been caught would produce the “phooey” and ugly face, she knew perfectly well. Being the self-confident individual she was, I have no doubt she hated any “phooey.” That I could see clearly from her expression on the few occasions, I had had to use it. She had been brought up to think for herself, to be imaginative and creative, and to believe in herself, not to be a pitiful dog waiting for her master’s voice before daring to blink.

If Petrine had rejected “phooey” as an unacceptable means of solving the conundrum, the only way to come out of it without losing face was to do what she did. She actually controlled the situation. If it is true that I could trigger her retrieving behavior (and that, combined with searching, was our best game in the whole wide world), by just assuming any position remotely resembling the game, so too could she trigger my behavior, my part in the game. That, she did indeed. She showed me a perfect retrieve, and put me in my role in the game. “Your line, now,” she said to me, clearly and emphatically without even the need of words. Like an experienced actor playing a Shakespearian part, I reacted promptly to my cue.

If a behavior repeated often with fairly predictable consequences creates moods (Pavlovian conditioning) in all of us, independently of species, which seems to be the case, I have no doubt that she associated the retrieve game with the most pleasure she could have in life. When in trouble, we have a tendency to perform behaviors that previously have brought us success, pleasure. This is a reassuring procedure, the basis even for stereotyped behaviors according to some. It is an organism’s attempt to re-establish emotional (neurophysiologic) homeostasis. If this is the case, Petrine’s solution was a good one, an intelligent one (as we would say of ourselves) and entirely compatible with our body of knowledge. It may seem improbable at first, but it becomes more reasonable the more we think about it.

Some of you will still think I am anthropomorphizing, and you have every right to do so. Pre-Petrine era, I would have thought the same. I would never have conceived of such an explanation. However, post-Petrine, a little dog that helped me discover many facets of life on Earth, I’m no longer so sure of the boundaries of anthropomorphism. Are intelligence, reasoning and self-respect only human features? In my opinion, as an evolutionary biologist, it is unlikely. Maybe language is misleading us once again. As Carl Sagan wrote, “Absence of evidence is not evidence of absence.” After all, why should “we” be so radically different from “them”?

Whilst I wouldn’t dare to rely on the unobservable self-respect on a scientific study, I wouldn’t dare either not to rely on it at a personal level on any one-on-one relationship independently of species involved. Unobservable and un-measurable, it may be, yet it remains for me a solid guideline reminding me that I am but one among many.

Life is great!

Featured image: Petrine, the English Cocker Spaniel, compelled me to ask: are intelligence, reasoning and self-respect only human features?

Odie The Pekinese: Awaiting On Death Row


Odie came to me on an odd day, one of those rainy, grey days, when the only thing you want to do is stay at home, listen to good music, watch the fire roaring in the fireplace, hold a hot cup of punch in your hands and feel sorry for yourself. Odie, an ugly duckling of a Pekinese, was awaiting his turn on death row. A twist of fate meant Odie survived his death sentence and, one year later, he had turned into a beautiful wolf.

I was sitting in my office at my desk, gazing absent-mindedly at a blank piece of paper lodged in my typewriter, which, unfortunately, had been stuck there for far too long. I was suddenly wrenched from my thoughts when our vet knocked at the door. “Have you got a minute? ” she asked. I debated saying “No,” but overcame the temptation. She came in, accompanied by Odie’s owners, and explained the situation. Odie’s owners wanted to euthanize him because they were sick of a particularly annoying behavior of his. He urinated all over the house and, when one day they found him cocking his leg up the impeccable flower arrangement they had proudly positioned in the middle of their much cherished, antique mahogany dining table, that was the last straw.

“Right on top of the table?” I asked them and they nodded solemnly.

I glanced down at Odie with newfound respect for it was no mean feat for an eight-inch (20 cm) tall Pekinese to climb on top of a dining table in order to accomplish a vital mission. So I asked them if I could keep the dog instead of them euthanizing him. I would try to solve his problem and find a good home for him. They were overjoyed at my proposal and I thus found myself being the improbable owner of a Pekinese for the first, and no doubt, last time in my life.

I was on a very tight deadline to write an article. After giving Odie a quick once over, I turned back to my typewriter and the embarrassingly blank sheet of paper. I remember thinking “Gee, you’re a really ugly little fellow, I understand why they wanted to get rid of you.” Odie grunted once in return. I think he could take a bit of humor. I would take care of Odie later. My first priority was to fill that all too white sheet of paper with some wise words.

Once deeply submerged in writing my article (or not writing it as the case may be), it was then I heard an almost imperceptible sound that took a couple of seconds to register and identify. I spun round to the source of the sound and, to my astonishment, my suspicion was confirmed. Odie was peeing on my books on my bookshelf.

I am a peaceful person and it takes a lot to upset me. Being a child of the sixties, I accept everyone and almost everything; all is good as long as it doesn’t restrict my freedom. However, one thing I must confess I can’t take is having someone peeing on my beloved books. I don’t discriminate: nobody urinates on my books, period! My reaction was therefore pure reflex. I reached for the first thing I had at hand, ironically enough it was my first book about dog training and behavior “Psychology Rather Than Force” and, before I knew it, I had thrown it at Odie.

The book, a good quality hardback, landed with a smack right behind Odie. Taken by surprise, he yelped, performed a beautiful pirouette in the air and stood there looking baffled and bewildered, staring at my book. For my part, I remained quiet as a mouse, holding my breath. After a few seconds, Odie managed to compose himself. He approached the book, sniffed at it in a noisy, Pekinese manner, then took a sniff at the bookshelf before returning to my book on the floor and giving it another long, even noisier sniff. Smacking his lips, he decided to lie down right next to the book. I returned to my tauntingly clean sheet of paper whilst keeping one eye on Odie.

Odie fell asleep, or so it seemed, and I finally began filling the blank sheet of paper with some meaningful words. A little later, whilst searching for something on my desk, I happened to knock a pencil over the edge and it fell on the floor between the desk and that bookshelf, a source of so much knowledge and inspiration. Odie opened his big, bulging eyes, one looking right and the other looking left, and approached the pencil. I couldn’t see him or the pencil but could hear him clearly, grunting, snuffling, puffing and panting. A few seconds later, maybe 15, he came around the desk directly towards me. He was holding the pencil in his mouth, each eye still looking in a different direction, one as wet as the other, dribble all over his face, with his head covered in balls of dust and fluff reminding me that my office needed a good hoovering.

I stretched out my hand to him and automatically said “tak” (which means “thanks” in the Scandinavian languages and was my sound signal for “release”). Odie, with a grunt, promptly dropped the slimy pencil into my hand. I was impressed. Was that a “retrieval”? Did he really retrieve that pencil for me?


Odie became very popular. His odd looks combined with his skills were an improbable combination in most people’s eyes.

I was so baffled and curious that I proceeded to do something that fellow pencil lovers regard as the ultimate sin towards pencils. You never drop a pencil as it is highly likely you’ll break the lead inside, rendering it useless once sharpened a couple of times. I tossed the pencil so it fell in the same place between my desk and the bookshelf; and once again, Odie ran (I think he was running, but don’t know for sure as I couldn’t see his short legs for all the fur), he grunted, snuffled, puffed and panted, rubbing one eye then the other along the floor in an effort to pick up the pencil and, in doing so, collected even more dust fluff. He wouldn’t give up, finally managed to take the pencil in his mouth and promptly returned it to me just as he had done before.

“Hallelujah!” I exclaimed despite my lack of religious conviction, “We have a retriever!” Joy filled my heart. The misery and self-pity the dull, grey day had imposed upon me ever since I had got out of bed that morning were gone like magic. Of all the activities I have undertaken with dogs, the one that has most amused me, and my dogs too it would seem, is without a shadow of doubt search and retrieve.

Odie never again urinated indoors, a fact we have discussed at some length. We are convinced it was the book incident that did it, due to the optimal coincidence of a series of conditions. Firstly, he was caught in the act (perfect timing), secondly, he did not associate the book falling behind him with me (instead with his own behavior), thirdly, the smack of the book falling on the floor had the right intensity to startle him (not too much, not too little), and fourthly, he associated the book aversive with his urinating behavior and nothing else (it happened when he urinated, it stopped when he stopped). No bad feelings towards books and (of course) no bad feelings from books towards him. Of course, the moral of this story is not that you should throw books at your dog. Let me say this loudly and clearly so no one gets it wrong: I do not recommend people throw books at their dogs. It worked in this case because of the coincidence of the many necessary conditions for it to work (as I explained) and that’s it.

I kept Odie and we all trained him. Sit, stand and, down were no problem at all, only difficult to observe for all the fur and short legs. We used treats as unconditioned reinforcers and my “dygtig” (as a semi-conditioned reinforcer), but he would do anything as long as we held a pencil in our hands (this was his reinforcer of choice). He would take the treats only because he was hungry. We put him on a program where he had to work for all his food and he worked a lot: no free food at all. Odie became very popular. His odd looks combined with his skills were an improbable combination in most people’s eyes. The staff at the Ethology Institute sometimes asked if they could take him home to show visiting friends. Odie never disappointed.

At the time, I was living in one of those enormous, old European mansions, like small castles, with three floors and endless of rooms. One particularly cold winter when the fields were covered by snow and ice, our cellar (basement) became a refuge for mice. This is very normal and we all know how to deal with the problem, except that I thought at the time it was more dignified for a mouse to die in battle than to be trapped or poisoned. Therefore, I introduced a hunting session every night at 8 pm after having read my son Daniel his bedtime story.

The nightly hunting session began with the troops, Petrine, Elanor (English Cocker Spaniels) and myself, assembling at the door to the cellar. Petrine and Elanor were skilled hunters so this was a good opportunity to stimulate them. Every evening we enacted the age-old game of predator and prey in the cellar of that big, old mansion house. Odie was always very keen to join us on our mission and, one evening, I decided to let him give it a go. Odie experienced his first hunt.

Odie quickly learned the rules of the game, although learn is perhaps the wrong term as it looked like he had always known and just had to be reminded. The first time, he went under a couch to chase a mouse, he took a long time. All I could hear was his usual grunting, snuffling, puffing, panting and the occasional high-pitched squeak from a mouse. I guess the mice were terrified of Odie’s looks combined with the spluttering, snorting and grunting. He came to me carrying his first mouse by a hind leg, the mouse completely stiff and wet, but very much alive. Odie became an efficient mouse hunter. He was quick and could squeeze into confined spaces for which the cockers were too big. Every evening, he was the first to reach our rendezvous point. He was there from around seven onwards, waiting patiently. He insisted on being the first to reach the bottom of the stairs to the cellar which was quite a spectacle for the steps were too steep for his all too short legs. He somehow managed to overtake the cockers on the way down, not running, but tumbling down amidst a cloud of dust and much snorting and grunting. The cockers just looked at him bemused. Up until then, our mission had been a well-planned military operation. Stealth, discipline, training, dedication and precise timing were our weapons. After Odie joined us, it all looked more like Asterix and Obelix against the Romans.

The days passed, one year passed, and Odie grew older and more experienced. I bet he could have won all kinds of competitions, but we never subjected him to that. By then he had become a great hunter, only limited by his physical characteristics, the ones us humans have bestowed upon him through selective breeding.

It was bound to happen sooner or later: one day someone came along that wanted to keep Odie. It was love at first sight when they saw his antics. When they asked me about his original problem, I couldn’t even remember what it was. I had completely forgotten, as had we all. After that first “attack” by my book, he had never again urinated indoors. Odie found a good home, one year after he had entered our lives.

I was sad to see him go. We all were. We often spoke fondly of him and made each other laugh by telling Odie stories. Odie had taught us invaluable lessons. First, that we should never judge anyone by their appearance. He was a little dog, short-legged, furry, flat-faced and cross-eyed, but he was a dog at heart like any other. None of us thought he was ugly, despite my initial horror. He was further evidence that many dogs develop problems because they are not treated as dogs; they are understimulated and their excess energy causes them to engage in any kind of activity, be it desirable or undesirable for the owners. He was a quick learner and an impeccable hunter with an enormous joy for life. Without words, he told us: “Respect and you shall be respected. I’m not a toy, not a thing, not a little human. I’m Odie, a Pekinese dog.”

14 years later, I went to give a talk in a town about 50 km from where I lived. During the break, a couple approached me and asked me if I remembered them. It took me a while, but I did recognize them. They were the new owners we had found for Odie. He was still alive, they informed me, but very old and tired by then. He no longer had any front teeth, as his love for retrieving hard objects had not waned over the years. They said they were getting ready for the day they would have to say goodbye to Odie and I saw their eyes well up.

Thinking of him, my eyes welled up too, Odie, the ugly duckling of a Pekinese that had turned into a beautiful wolf in my eyes and in the eyes of all those who had the privilege to know him. Thanks, Odie, my friend!


PS—I know that metamorphosis does not occur in canids and that a dog cannot turn into a wolf. I also know that a dog is a dog (Canis lupus familiaris) and not a wolf (Canis lupus lupus). Since this is a story with a point written for a blog, not a scientific article, I allow myself some artistic license when I write “Odie turned into a beautiful wolf.”

Featured image: Odie, an ugly duckling of a Pekinese, was awaiting his turn on death row. A twist of fate meant Odie survived his death sentence and, one year later, he had turned into a beautiful wolf (Photo by Nikolaj).

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Cultural Differences in Dogs—Our Responsibility

Cultural Differences Dogs

Do animals have cultural differences? Do dogs behave differently according to their doggy culture?

Yes, they do. Normal behavior is only normal under specific conditions. Normal behavior is the behavior displayed by the majority of the population in a precise area in a particular period. We may not like it, but if most do it, then it is normal. An extreme example: to behave rationally is not normal among humans since most people behave irrationally.

Yes, dogs show cultural differences. Their facial expressions and body languages show slightly different nuances from region to region. Even barking and howling can be distinctive. Davis Mech discovered that when he flew to the Abruzzi Mountains in Italy to assist Luigi Boitani and Erik Zimen with their wolf research. The Italian wolves howled with an accent (or so did the Americans).

Natural selection determines the cultural differences our dogs show from one area to the other. We breed those we like best, and we like them differently from place to place. Remember, selection acts upon the phenotype (the way a dog looks and behaves), but the traits pass to the next generation thru the genes (genotypes) involved in the favored phenotypes. Don’t forget as well that our human choices as to preferred animals are also natural selection.

Cultural differences in dogs are easy to spot when one travels as much around the world as I do. It still surprises me, for example, to see that European English Cocker Spaniels or Bichon Havaneses are so very different from their American counterparts—and not only physically, also behaviorally—same breeds, different cultures.

A culture develops according to the influence of the particular individuals in a group and their distinct environment. The unique characteristics of the individuals and the environment determine cultural development. In dogs, we are the most influential environmental factor. Therefore, the same original breeds develop variants depending on the human group with which they interact. Undoubtedly, we create the various canine cultures we have. The question is whether we do it intentionally or unintentionally, and I believe we can argue for both.

In a sense, we can say we have the dogs we deserve. We have created them, either by planning to breed them to achieve specific results or not caring at all—which amounts to the same in this context. The problem is that cultures evolve, our societies change, and so do our needs and requirements—which is fair enough. What does not seem fair to me is to impose new (cultural) requirements upon the dog all of a sudden after having created it to fulfill different (cultural) needs back in the day. Of course, we (as a species) change with time, and so do our dogs. We can change them, the dogs, to meet new necessities, as we’ve done in the past, but it requires a well-planned breeding program based on reasonable expectations and scientifically sound methods. 

The ethics of changing animals as we please is an entirely different discussion, alas one for another day.




Cavalli-Sforza, Luigi L (1986). “Cultural Evolution”American Zoologist26 (3): 845–855. doi:10.1093/icb/26.3.845.

Davis, N.B.; J.R. Krebs, J.R.; West, S.A. (2012). An Introduction to Behavioural Ecology (4th ed.). ISBN-10:9781405114165.

De Waal, Frans. (2001). The Ape and the Sushi Master: Cultural Reflections by a Primatologist. New York: Basic Books.

Holdcroft, D.; Lewis, H. (2000). “Memes, Minds, and Evolution.” Philosophy75 (292): 161–182.

Jones, Nick A. R.; Rendell, Luke (2018). “Cultural Transmission.” Encyclopedia of Animal Cognition and Behavior. Springer, Cham. pp. 1–9. doi:10.1007/978-3-319-47829-6_1885-1. ISBN 978-3-319-47829-6.

Laland, Kevin N. and Bennett G. Galef, eds. (2009). The Question of Animal Culture. Cambridge, Massachusetts: Harvard UP.

Swanson, H.A., Lienand, M.E., Ween, G.B. (2018). Domestication Gone Wild: Politics and Practices of Multispecies Relations. Duke University Press.

Featured image: Dog behavior shows cultural differences across breeds and regions (photo by NewEvolution at

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Learn more in our course Ethology. Ethology studies the behavior of animals in their natural environment. It is fundamental knowledge for the dedicated student of animal behavior as well as for any competent animal trainer. Roger Abrantes wrote the textbook included in the online course as a beautiful flip page book. Learn ethology from a leading ethologist.

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Fearful Behavior—Genetics and the Environment

Fearful Dog (Fearful Behavior—Genetics and the Environment)

Two essential aspects of fear: (1) Fearful behavior has genetic and learned components, and (2) our pets may show fearful behavior because we have taught them that without being aware.

We usually distinguish between rational or appropriate and irrational or inappropriate fears. The latter are called phobias, i.e., fears that are disproportional to the dangers in question, although some phobias do have a survival value.

Fear mechanisms serve the survival of organisms by producing appropriate behavioral responses. Hence, evolution has preserved it, subject to adaptive changes throughout time and according to the posed environmental challenges. From an evolutionary perspective, the particular fear behaviors of a species may be an adaptation that was useful at some point in the past. The distinctive responses to fear stimuli may have emerged and developed during different periods. For example, fear of heights, common to most mammals, has probably developed during the Mesozoic period; and fear of snakes, usually in simians, during the Cenozoic period. Claustrophobia, agoraphobia, and aquaphobia may also have their origins in evolutionary adaptations.

Predators and prey have different strategies to deal with threats. Their behavioral strategies evolved throughout millennia under the constant struggle for survival. Predators avoid dangerous stimuli by creating distance, escape being the favored strategy. Prey animals freeze preferentially when the predator is still relatively far away, but when distance decreases to a critical value, the animal flees. Thus, flight appears to be a genuine unconditional response to the unconditional stimulus consisting of a predator at a critical distance.

Fearful responses and their intensity seem to be a consequence of predisposing traits, resulting from many gene-environment interactions during the development of the individual. The latest research has established a genetic basis for fearful behavior. Researchers conducted studies with humans as well as other animals. In humans, researchers have been able to study the effect of genetics (family lines and twins) and environment (adoption cases).


Fleeing is the first strategy when facing a threat. Horses seldom gallop in nature except when fleeing from a predator.

Flight is the primary strategy animals use in the face of a threat. This behavior is associated mainly with the sympathetic nervous system, which mobilizes the organism for the so-called fight-or-flight response originally described by Cannon. While flight is an active coping strategy when facing danger, freezing (immobilization) and hiding are passive coping strategies. Depending on species, such a strategy is the next-best option to flight. Animals freeze and hide when escape is impossible. Freezing implies an inhibitory activity in the autonomic system (hypotension, bradycardia), formerly described by Engel and Schmale as a conservation-withdrawal strategy.

Thanatosis, or tonic immobility, is an extreme form of freezing behavior. For example, white-tailed deer fawns (Odocoileus virginianus) can lower their heart rate to 38 beats per minute (from about 155) for up to two minutes.

Whether an animal has a preference for an active or a passive defense strategy is not solely a question of context. Research shows that some animals do prefer one strategy rather than the other. In exactly the same situation, two animals may respond differently. The interesting point is that these patterns, both behavioral and neuro-endocrinal, seem to be consistent. Some researchers suggest that this may explain why some individuals are more resistant to stress and stress-induced malfunctions than others. Researchers found the tendency to react one way rather than the other to run in families, suggesting a genetic component. The experiments were conducted with rats and mice, but we have no reason to suspect that studies of other species would not yield the same results.

Fear is probably experienced similarly in many species. All mammalian species show three different sites in the brain where electrical stimulation will produce a complete fear response: (1) the lateral and central regions of the amygdala, (2) the anterior and medial hypothalamus, and (3) areas of the PAG, the periaqueductal gray, which is the gray matter in the midbrain involved in the modulation of pain and defensive behavior. Researchers have also studied defensive strategies in various species and concluded that human reactions to threatening stimuli are not qualitatively different from nonhuman mammals.


The early development has a critical influence on how animals will respond to challenges, stress and fear eliciting stimuli.

The amygdala seems particularly relevant. We suspect that it may have a significant function in regulating many facets of social behavior. It also appears that threatening stimuli activate the amygdala, which in turn has a decisive influence on the cognitive mechanisms of the individual, including the perception of the environment, selective attention (relevant for learning), and memory.

Conclusion: Not surprisingly, and in line with many other behavioral traits, fearful behavior depends upon two different factors: (1) a genetic predisposition and (2) the influence of the environment. Environmental factors during the development of the young individual may be critical in its ability to cope with stress and fear-eliciting stimuli. Early experiences appear to affect the neural and biochemical systems involved in fearful behavior and in dealing with stress—as well as learning processes and the capacity to deal with threatening stimuli in adulthood. Maternal prenatal stress may also produce changes in the brain morphology of the fetus and, consequently, in its way of reacting to stress and fear-eliciting stimuli, later in life.

While some fear responses are innate, others are learned. Conditional fear provides a critical survival-related function in the face of a threat by activating a range of protective (or defensive) behaviors. Therefore, we can presume that all animals will be ready to identify and retain the memory of any stimulus or situation they have perceived as potentially dangerous or threatening. Thus, it is natural and easy for animals to develop fearful behavior.

Watson demonstrated how fear could be a conditioned response with his famous (or infamous) experiments on Little Albert in 1920, who learned to fear a white rat. Some of the fear behavior of our pets, particularly dogs and cats, are created by us. An event that in itself might pass nearly unnoticed may be blown up to a disproportionate relevance if associated with a strong reaction of the owner. Dogs (and children) often face situations with unexpected and somehow aversive results, which they would soon forget if it weren’t for the exaggerated reaction of the owners (parents). All living organisms are, in principle, prepared to deal with discomfort, aversive experiences, and failure. The problem is when these assume proportions out of context because they are additionally reinforced. For example, many dogs fear strangers because their owners fear that the dogs fear strangers, and their reactions reinforce the dogs’ disposition to be cautious about strangers. Often, and unaware of it, the owner reinforces the fearful response while attempting and believing that he/she is reassuring the dog. That is conditioned (learned) fear behavior.

We saw it clearly in the 1980s when we performed some experiments at Ethology Institute. A litter of puppies from a suspected line of dogs prone to show fearful behavior exhibited entirely distinct behaviors one year after we had placed them in six different homes. The dogs reflected, indeed in a significant degree, the attitude of their owners toward novelty and challenges. We repeated the experiment with another litter, this time from a confirmed non-fearful line, and the eight puppies showed the same tendency again when we tested them one year later. Even though there was a tendency for the dogs from the fearful line to be on average more cautious and the others to be bolder, they overlapped one another in the middle range of responses. Our tests did not include enough animals to enable us to draw a conclusive answer as to the question of genetics versus the environment in this aspect. However, they pointed out the importance of the environment, at least in what concerns the average domestic setting in which we can expect dogs to grow.


Cannon, W. B. (1915). Bodily Changes in Pain, Hunger, Fear and Rage. New York, NY: Appleton.

Engel, G. L. and Schmale, A. H. (1972). Conservation withdrawal: a primary regulatory process for organic homeostasis. In: Physiology, Emotions and Psychosomatic Illness. New York, NY: Elsevier; 1972:57–95.

Kavaliers, M. and Choleris, E. (2001). Antipredator responses and defensive behavior: ecological and ethological approaches for the neurosciences. Neurosci Biobehav Rev. 2001;25:577–586.

Koolhaas, J. M. et al. (1999). Coping styles in animals: current status in behavior and stress-physiology. Neurosci Biobehav Rev. 1999;23:925–935.

McFarland D. (1987). The Oxford Companion to Animal Behaviour. Oxford, UK: Oxford University Press.

Panksepp, J. (1998). The sources of fear and anxiety in the brain. In: Panksepp J, ed. Affective Neuroscience.New York, NY: Oxford University Press; 1998:206–222.

Parmigiani, S., Palanza, P., Rodgers J. and Ferrari, P. F. (1999). Selection, evolution of behavior and animal models in behavioral neuroscience. Neurosci Biobehav Rev. 1999;23:957–970.

Perrez, M. and Reichert, M. (1992). Stress, Coping, and Health. Seattle, Wash: Hogrefe & Huber Publishers.

Steimer, T. (2002). The biology of fear- and anxiety-related behaviors. Dialogues Clin Neurosci. Sep 2002; 4(3): 231–249.

Watson, J. B. (1970). Behaviorism. 7th ed. New York, NY: WW Norton & Company.

Weinstock, M. (2001). Alterations induced by gestational stress in brain morphology and behaviour of the offspring. Prog Neurobiol. 2001; 65:427–451.

Featured image: Dog showing fearful behavior. Paw lifting indicates a beginning of pacifying behavior (photo by Lifeonwhite).

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Learn more in our course Ethology. Ethology studies the behavior of animals in their natural environment. It is fundamental knowledge for the dedicated student of animal behavior as well as for any competent animal trainer. Roger Abrantes wrote the textbook included in the online course as a beautiful flip page book. Learn ethology from a leading ethologist.

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Fearful Behavior—the Making of a Definition

Fearful Behavior (The Making of a Definition)

We have discussed aggressive behavior (1 and 2). We shall now examine fearful behavior beginning (as always) with a definition, but first, let us look at existing definitions.

Fear is an unpleasant often strong emotion caused by anticipation or awareness of danger (Merriam-Webster).” As a dictionary definition (and that’s what it is), it works, but we need to be more precise.

Fear is the unpleasant emotional state consisting of psychological and psychophysiological responses to a real external threat or danger.” (Miller-Keane Encyclopedia and Dictionary of Medicine, Nursing, and Allied Health, Seventh Edition). This definition is more precise than the former. However, to evaluate it correctly, we need first to define threat, particularly as to what real external threats are versus what the text leaves us presuming are not real threats. Second, the term unpleasant is vague. How can we observe psychological responses? We cannot. What we observe are behavioral changes, but that is not what the definition says. We can measure some psychophysiological responses, but I wouldn’t rely entirely and solely on them to analyze the consequences of a frightening experience.


When facing a threat, and flight is not possible, the next best option is to hide and freeze (photo from Molly’ s Just You Only Better).

“Fear is a normal emotional response to consciously recognized external sources of danger such as those often associated with loud noises, threatening gestures, strange people and thunderstorms; it is manifested in animals by flight, by attack or by cringing.” (Saunders Comprehensive Veterinary Dictionary, 3 ed.). This definition contains the explanation of real threats (consciously recognized external sources of danger) that we missed above. It also gives us examples. Whether fear is manifested by flight, by attack or by cringing is controversial for an ethologist because these all have different functions. They appear together is this definition, probably because veterinarian science tends to classify behavior by symptoms (as is the practice in its field) and not function as evolutionary biologists (and ethologists) do. A fearful stimulus may trigger an attack (function=eliminate a threat), but an attack is aggressive behavior, even when defensive (function=eliminate competition), and cringing (a term not commonly used in the behavioral sciences) may signify submissive behavior, which is not the same as fearful behavior (function=eliminate a social threat).

“Fear is an emotion induced by a threat perceived by living entities, which causes a change in brain and organ function and ultimately a change in behavior, such as running away, hiding or freezing from traumatic events.” (Wikipedia). That is a good definition. It relates emotion with behavior, which is always sweet music to the ethologist’s ears, who prefers to deal with observable and measurable phenomena rather than the occult emotions. Running away, hiding or freezing are compatible as to function, so we have no problem with that. Traumatic events, on the other hand, requires an explanation.

Emotions are experienced and expressed at three different levels: (1) the psychological level, (2) the neurophysiological level, and (3) the behavioral level. All three aspects are present in all emotions. While psychologists focus on the first level and psychiatrists on the second, ethologists concentrate on the third.

“The main function of fear and anxiety is to act as a signal of danger, threat, or motivational conflict, and to trigger appropriate adaptive responses. For some authors, fear and anxiety are indistinguishable, whereas others believe that they are distinct phenomena.” (Steimer). That is a short, precise, and strong definition. It gives us the function of fear and points out an important distinction to a related term, anxiety, which “[…] is a generalized response to an unknown threat or internal conflict, whereas fear is focused on known external danger.”

Steimer’s and my definition are fully compatible. Here is mine:

Fearful behavior is behavior directed toward the elimination of an incoming threat, e.g., fleeing, freezing, or hiding. Submissive behavior is behavior directed toward the elimination of a social-threat from a mate, i.e., losing temporary access to a resource without incurring injury, e.g., and highly depending on species, lying down on the back, assuming a low-profile body posture, or turning the neck away.”

My definition is identical to Steimer’s, only focusing on the behavior and compiling all triggering factors under one label, threats. Moreover, I prefer to pinpoint the distinction to the related submissive behavior (instead of anxiety). Mine is a more extreme ethological definition and has the advantage of adding an explanation of submissive behavior. Steiner’s has the advantage of relating to anxiety and is more likely to be adopted by human psychologists and psychiatrists.

Both Steimer’s and my definition require a definition of threat. Steimer does not explicitly give us one. However, I do, and one, which is compatible with both our definitions.

“A threat is everything that may harm, inflict pain or injury, or decrease an individual’s chance of survival. A social-threat is everything that may result in the temporary loss of a resource and may cause submissive behavior or flight, without the submissive individual incurring injury.”


When facing a social-threat, submissive behavior is the best option. This dog shows what ethologists call active-submissive behavior (photo from safekidssafedogs).

I’m compelled (Steimer is not) to distinguish between threat and social-threat because, in my definition, I pinpointed the difference between fearful and submissive behavior.

I have used a term that also needs a definition, namely, mate.

“Mates are two or more animals that live close together and depend on one another for survival. Aliens are two or more animals that do not live close together and do not depend on one another for survival.”

McFarland explains fear as a motivational state triggered by particular stimuli that give rise to defensive behavior or escape. Fear is one of the primary motivators because of its life-saving function. Newborns and infants show innate fear responses to particular stimuli that may harm them. Also, environmental disturbances require habituation. Their fear of novelty loses its strength gradually and only comes back much later when trying new ways amounts to the spending of more energy (heavily penalized in nature) than doing it as they always have done.

Fear sometimes occurs in conjunction with other motivators. Approach-escape conflicts are typical examples and often result in displacement behavior as, for instance, self-grooming. We may argue that displacement behavior indicates the kind of fear that psychologists call anxiety. However, ethologists need not introduce a new term because the definition of fear includes the biological aspects of anxiety.

Even if anxiety and fear are probably distinct emotional states, there may be some common aspects in their brain and behavioral mechanisms. As Barlow says, anxiety may just be a more elaborate form of fear enabling the individual with an increased capacity to adapt and plan for the future. In Steimer’s words, “If this is the case, we can expect that part of the fear-mediating mechanisms elaborated during evolution to protect the individual from an immediate danger have been somehow ‘recycled’ to develop the sophisticated systems required to protect us from more distant or virtual threats.”

When describing animal behavior, it is more useful to refer to fear and fearful behavior than anxiety. Additionally, we must bear in mind that fearful is an adjective of behavior. To label a particular animal as fearful might be to go over the top. An animal may show fearful behavior, and rightly so, in certain situations and not in others.

Fear and fearful behavior evolved with the vital function of protecting the individual and are, therefore, mechanisms, which we may presume to have a strong genetic correlation. Animals of different species show fearful behavior to different stimuli and in different degrees. It is natural and normal for almost all animals (if not all) to get startled by a loud noise or a sudden movement. Horses and Guinea pigs get startled by more and different stimuli than dogs because they are prey animals, but they come over it quickly. They are not more fearful animals per se—they are just different, and comparison at this level is meaningless.

Two essential aspects of fear: (1) Fearful behavior has genetic and learned components, and (2) not seldom, our pets show fearful behavior because we have taught them that without being aware. These are the topics for the following article.




  • Barlow, D. H. (2000). Unraveling the mysteries of anxiety and its disorders from the perspective of emotion theory. Am Psychol. 55:1247–1263.
  • LeDoux, J. (1998). The Emotional Brain. New York, NY: Simon & Schuster.
  • McFarland D. (1987). The Oxford Companion to Animal Behaviour. Oxford, UK: Oxford University Press.
  • Steimer, T. (2002). The biology of fear- and anxiety-related behaviors. Dialogues Clin Neurosci. Sep 2002; 4(3): 231–249.
  • Strongman, K. T. (1996). The Psychology of Emotion. Theories of Emotion in Perspective. Chichester, UK: John Wiley & sons.
  • Watson, J. B. (1970). Behaviorism. 7th ed. New York, NY: WW Norton & Company.

Featured picture: Fearful and submissive behavior overlap. Fearful behavior is always submissive but it makes sense to distinguish between the two because some submissive behavior is not particularly fearful (rather respectful, in human terms). This excellent photo by Monty Sloan shows this overlap of motivational factors. The highlighted wolf shows more submissive and less fearful behavior than expected, given the clear aggressive and dominant display of its opponent. That is a sure indicator that they are mates, i.e., belong to the same pack.

Learn more in our course Ethology. Ethology studies the behavior of animals in their natural environment. It is fundamental knowledge for the dedicated student of animal behavior as well as for any competent animal trainer. Roger Abrantes wrote the textbook included in the online course as a beautiful flip page book. Learn ethology from a leading ethologist.

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Aggressive Behavior—Inheritance and Environment


This dog shows self-confident (dominant) aggressive behavior. This is instrumental aggressive behavior (photo from dog-adoption-and-training-guide).


This dog shows insecurity and aggressive behavior. This may be reactive-impulsive aggressive behavior, but may also be learned behavior (photo by


This dog (to the right) shows learned aggressive behavior. It may be impulsive-reactive, but it does not need to be (photo by

Having dealt with the definition of aggressive behavior in an earlier article, we will now analyze the various types of aggressive behavior and their correlation to genetics. Although a strong definition of aggressive behavior is a promising step to understand it, we have not resolved all matters and still need to clarify a few other terms. Note that in the following, to make it shorter, we will use aggression and aggressive behavior interchangeably. 

When studying human aggression, it is common to subdivide it into two types: (1) instrumental aggression, which is purposeful or goal-oriented; and (2) reactive-impulsive aggression, which is elicited by loss of emotional control and often manifests itself as uncontrollable or inadequate actions.

Let us also note that, “Aggression differs from what is commonly called assertiveness, although the terms are often used interchangeably among lay people (as in phrases such as ‘an aggressive salesperson’” in Quadri and Vidhate’s words. They also state, “Predatory or defensive behavior between members of different species may not be considered aggression in the same sense.” I would go a step further and insist we do not consider these behaviors as aggressive behavior in any sense.

Aggression Types

A distinction in types of aggressive behaviors is between (1) pro-active (also controlled and instrumental) and (2) reactive-impulsive. The former is not an end in itself, only the means to achieve a goal. There are no strong emotions involved. On the contrary, its effects depend on deliberate and well-timed action. The latter has no goal in itself and is marked by intense emotions. In short, researchers of aggressive behavior in children have found it helpful to distinguish between reactive (impulsive) from proactive (instrumental) aggression.

Modern frustration-aggression theory claims that anger is a reaction to an aversive experience, including frustration. It emphasizes the importance of moral violation as justifying the expression of aggressive behavior.

The question is whether we also find these types of aggressive behavior in animals other than Homo sapiens sapiens. Being an evolutionary biologist and a good Darwinist, I am always highly suspicious of any statement claiming that a trait is exclusive to one single species. The odds of that happening are worse than winning the big lottery.


Do animals other than humans have morality, and will they fight for a cause? That is a difficult question because I cannot envisage any way of verifying it. In that sense, some would even call it a meaningless question. 

Let us analyze the evidence we have. We know that some animals show empathy and altruism, widely recognized as conditions for morality. Shermer points out that humans and other social animals share the following characteristics: “[…] attachment and bonding, cooperation and mutual aid, sympathy and empathy, direct and indirect reciprocity, altruism and reciprocal altruism, conflict resolution and peacemaking, deception and deception detection, community concern and caring about what others think about you, and awareness of and response to the social rules of the group.”

However, we can account for all these characteristics in terms of evolutionary costs and benefits and using models based on evolutionarily stable strategies. We need not introduce a new term, morality, to explain that. Therefore: if humans show moral behavior, so do other species, albeit differently. What we might need to concede is that sometimes quantitative differences amount to qualitative differences. Hence, showing these traits to such a high degree, as is the case in humans, justifies us coining a new term, morality.

Suppose that is the case (and I’m only theorizing). Then, it makes sense to label some human behavior as moral and disregard the possibility of morality in other animals (unless remarkable discoveries enlighten us differently).

Thus, if it does not make sense to analyze non-human animals’ behavior in terms of morality, then it follows that we can neglect reactive-impulsive aggression caused by violation of moral rules in those animals.

However, we cannot dismiss the same behavior caused by loss of emotional control because non-human animals can also lose control over their emotions. The tricky part here is, as always, the term emotion, which is vague and, therefore, one that biologists prefer to avoid.



Emotions and Reactive-Impulsive Aggressive Behavior

What is an emotion? According to Schacter, an emotion is “[…] a positive or negative experience that is associated with a particular pattern of physiological activity” caused by hormones, neurotransmitters, dopamine, noradrenaline, serotonin, and GABA. We find all these in some non-human animals; therefore, if we can accept the above definition of emotion, we must concede that if we can show emotional behavior, so can they.

The only way it makes sense when dog people speak of dogs being reactive (meaning they growl, snarl, attack, or bite someone) is that dogs display reactive-impulsive aggressive behavior. It is still, by all means, aggressive behavior, just one type that may or may not exist to some considerable degree in non-human animals, depending on whether I am right or wrong in my theorizing.

Recognizing and identifying reactive-impulsive aggression may be advantageous because research shows that it may be easier deterred than instrumental aggressive behavior. Reactive-impulsive aggression appears to result from a distorted perception of competition, the aggressive individual not realizing that there are evading routes, and enhanced by the inability to control the associated emotions. There is also evidence that reactive-impulsive aggression (contrary to instrumental aggression) is related to low serotonin levels in the brain. On the other hand, classifying all canine aggressive behavior as reactive-impulsive, as it seems to be the practice these days, may turn into a fatal mistake with extremely severe consequences.

A dog displaying aggressive behavior can show it self-confidently (what we, ethologists, call dominant behavior) or insecurely (showing submissive behavior—not fearful). The former is not reactive-impulsive—it is instrumental and goal-oriented. The latter might be if the dog does not realize that a clear display of submissive behavior or flight would solve the problem. This kind of aggressive behavior may be: (1) the consequence of inadequate imprinting and socialization (the dog did not learn how to solve social conflicts), (2) the result of inadvertently reinforced behavior. Dog owners reinforce their dog’s reactive-impulsive aggressive behavior attempting to do what they call ‘calming down the dog.’ The dog growls, they say, “quiet ” (or similar), the dog looks at them, and they reinforce that with a treat and a “good job.” It doesn’t take many repetitions for the dog to learn that displaying aggressive behavior provides attention and food.

The term reactive does not belong to ethology, which classifies behavior by function. I don’t know how it came into dog training, but I suspect a psychologist introduced it, and dog people liked it because it sounded better to say, “My dog is reactive” than “My dog shows aggressive behavior.” Ironically, the term places the full responsibility for unwanted behavior on the owners—reactive-impulsive aggression is either the result of poor imprinting/socialization or inadequate training.


Is Aggressiveness Inherited?

Heritability studies attempt to determine whether a trait passes from parent to offspring. Some genetic lines in many birds, dogs, fish, and mice seem to be more prone to aggression than others. Through selective breeding, we can create animals with a tendency to show more aggressive behavior.

Some aggressive behavior is evolutionarily advantageous, and some are not and might be an impediment to social cohesion. Maynard Smith states that it is not surprising for aggressiveness to have a strong genetic correlation, given the high likelihood of both potentially positive and negative selective discrimination throughout evolution.

Research has uncovered many factors that contribute to aggressive behavior. Disruption of the serotonin system is a highly significant feature in predisposing aggression. There is a correlation between testosterone levels and aggression. Extremely low levels of blood sugar (hypoglycemia) may elicit physiological changes and aggressive behavior.

Most researchers agree that we must consider the influence of genes, not in isolation, but as functioning in the whole genotype, as well as the effect of the environment. Thus, future research in the genetics of aggressive behavior may very well focus on epigenetic factors.

Doubtless, most behavior traits, except simple reflexes, contain a genetic plus an environmental component. No behavior will develop without the appropriate genetic blueprint, and no behavior (again except for a few simple patterns) will show in the absence of the correct environmental stimuli.

It is probable each organism filters and selects stimuli from a wide range in its habitat according to its genetics, thereby creating its uniqueness of experiences. As Bock and colleagues say, we make our own environment. I have no reason to suspect that the same does not happen with other animals.





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Featured image: These two dogs are both equally self-confident (showing equal dominant behavior). Any aggressive behavior deriving from this situation will not be reactive-impulsive, but instrumental (photo from dog

Learn more in our course Agonistic BehaviorAgonistic Behavior is all forms of aggression, threat, fear, pacifying behavior, fight or flight, arising from confrontations between individuals of the same species. This course gives you the scientific definitions and facts.

Agonistic Behavior is a brand-new, for 2019 created course by leading ethologist Roger Abrantes. It is one of our most challenging courses, addressed to the dedicated student of behavioral sciences.


Love Your Dog the Whole Year

The big summer vacation is setting in all over most of the western world. Children have finished school for this (school) year. It’s a good, but also a stressful period.

At the Institute, every year, our IT department experiences its ‘low season’ in June-July and again in December. For us, IT does not stand for Information Technology, but Individual Training. Our activity of treating problem behavior individually predates the upsurge of computers and their IT acronym.

June-July is the big summer holiday season. Those who stay home have more time to give to their pets, which is good, but can also create problem behavior if we are not careful. Having more time at their disposition means that most dog owners spend more time with their dogs—more and longer walks, maybe a bit more training, but most of all much more time together.

All organisms are more or less sensitive to routine changes, and our dogs are not exceptions. The increase in activity with the children at home can imply some stress, and many dogs do not respond well to that. Some of them become restless, hyperactive.

Another problem occurs with being home alone, which our statistics shows clearly. If June-July and December are low seasons, then August and January are high seasons for canine problem behavior. Having been together with their owners almost 24-7 for a longer period, many dogs react poorly when the school and work routines set in again. From one day to the other, without any explanation (for them) all the razzmatazz disappears, and they are left home alone. They have been extremely busy for a period, and unexpectedly nothing happens around them.

From having company around the clock to being alone eight hours a day, or from being active most of the day to suddenly having nothing to do, creates problems.


Summer holiday—a great time for the whole family, including the dog. However, we must be attentive to problems we may be creating without being aware of it.

What can we do to prevent our dogs from developing behavior problems during the long holidays?

The time we spend with our dogs should be quality time more than quantity time (I guess this applies to all relationships). More time during the long holidays is good because we can focus on training our dogs in some skills, which maybe we’ve wished to do, but couldn’t because of our busy working schedule.

To prevent home alone problems from showing up after the holidays, we should maintain periods during the day in which the dog is unattended. This will also prevent the worst of the hyperactive behavior that may develop due to the higher level of stimulation. Instruct the children to leave the dog alone at set times and explain to them why this is important.

As to the activities, you can do with your dog, try to avoid the empty, stressing activities like ball throwing and chasing. Focus on the more meanings full searching games instead—nose work or scent detection as you prefer to call them. These are activities that tire the dog without creating hyperactive behavior. You can even teach your children these searching games. They will spend some good times with their dogs developing a healthy relationship. It is my experience that children are great with animals if we, adults, give it the necessary time to instruct them correctly.

If you go abroad on holidays, remember that boarding a dog is stressful for the dog independently of the quality of the boarding venue. The best of them will provide suitable conditions for the dog to satisfy its needs for contact and exercise, but it is still a break in the usual home routine. Be prepared to reintroduce the homely routines when you return from what I hope has been an enjoyable holiday trip to an inspiring foreign country. It does not need to be difficult if you are aware of it and do it systematically.

Other pets than dogs are also affected by our long holidays. Cats may become extremely restless during the holidays and seek refuge. Horses may show stereotypies when returning to the pre long holiday routine, if you have spent much more time with them. Even parrots have shown some problem behavior because of the disruption of the daily schedule.

It’s curious how our minds focus on irrelevant contexts and forget essential ones. For example, no canine has evolved to expect food presented at set times, and yet most dog owners insist in serving their dogs the daily rations by the clock. On the other hand, no canine has evolved to see their family-pack increase/decrease their level of activity dramatically from one day to the other (like with holidays) or to be moved suddenly to a totally unfamiliar location with a new fauna, and yet many dog owners don’t even give it a thought.

Our responsibility toward our dog is a whole year activity (as it is to be a parent). To love those we are responsible for means to provide them with what they need so that they can develop harmoniously, and, hopefully, create their own happy life. To love is a full-time job with no time off allowed.

Please, share this link with your clients, if you’re a dog trainer, and with your dog owner friends. If it only can help a few, preventing their dogs to develop problem behavior, then it’s worth it. It would be wonderful if we’d go out of business in what concerns treating problem behavior in pets, wouldn’t it? Alas, I don’t think that to be a realistic expectation anytime soon, but we can all do our part to help it move in the right direction.

Enjoy your holiday and keep smiling. Yes, life is great!

Featured image: Love is a full-time job (photo by Lauren Grabelle).