In my decades of work as a cynologist, I have encountered thousands of dogs and their owners across the world, from the precision-driven Belgian Malinois bred for elite police and military tasks, to the tireless Siberian Husky bred for endless Arctic miles, to the obsessive heelers and Australian Shepherds whose very DNA burns with herding intent. And in those decades, one pattern has repeated itself with such frequency that it has become nearly predictable:
An owner walks into my training center, exhausted, often embarrassed, sometimes even angry, and almost always overwhelmed. The story unfolds in variations, but the refrain remains the same: “Bart, this dog is out of control. He barks constantly. He jumps on people. He chews through furniture. He can’t settle down. Can’t you just train him to calm down?”
This plea is understandable. These owners are not bad people. Most of them truly love their dogs and want what is best. But their expectations, shaped by either dishonest breeders or cultural misunderstandings about what “companionship” means in a dog, are tragically mismatched with the biological reality of the animal they now live with.
The root of the misunderstanding lies in one crucial truth that must be stated plainly:
Drive is not a behavior. Drive is biology.
It cannot be erased by obedience commands, nor extinguished by stricter rules, nor suppressed ethically through training. Drive is the living current of motivation flowing through a dog’s neurobiology, carved by millennia of evolutionary selection and hundreds of years of targeted human breeding. To attempt to “train it out” is not only futile—it is, in most cases, cruelty disguised as obedience.
In ethology, the study of animal behavior, “drive” is often used to describe the motivational systems that propel an animal to engage in species-typical behaviors. In wolves, these drives include hunting, territorial defense, reproduction, and social cohesion. When humans domesticated and later selectively bred dogs, we did not eliminate these drives. Instead, we sculpted them to meet our utilitarian needs.
In some breeds, we amplified prey drive but truncated the sequence before the kill, producing herding dogs that stalk, eye, and chase but rarely deliver the final bite.
In others, we fortified defense drive, cultivating animals predisposed to stand their ground, bark at intruders, and fight if necessary.
In still others, like sled dogs, we maximized locomotor drive—the need to run, sustained not by immediate reward but by the neurochemical reinforcement of endurance itself.
Thus, the dog you live with is not merely an animal responding to training cues. It is the bearer of genetically sculpted motivational systems that exist independent of your household expectations.
The Persistence of Drive in Modern Society
The mismatch between dogs’ genetic drives and human lifestyles is one of the most pressing welfare problems of our time. A Belgian Malinois is not born understanding that he now lives in a suburban home where his task is to “just relax.” His neurology does not receive the memo. His dopamine system still anticipates the bite sleeve, the chase, the confrontation.
The same is true for the Australian Shepherd bought by a family for its “cute looks” or the Husky purchased because someone saw a viral video of one “talking.” These dogs do not transform into calm companions by virtue of living in a city apartment. The drives remain, and without appropriate outlets, they manifest as what owners perceive as “problem behaviors”: barking, destructiveness, hyper-arousal, reactivity.
But from the dog’s perspective, these are not problems. They are expressions of unmet biological needs.
Breed-Specific Expressions of Drive
Let us look more closely at how drives manifest differently across breeds.
A. German Shepherd Dog (GSD)
The Working line German Shepherd is perhaps the quintessential example of a high-drive working dog. Originally bred in late 19th-century Germany for herding, the breed was selected not just for stamina and intelligence but for the ability to make rapid decisions under stress. When herding tasks diminished, the GSD’s versatility made it a natural candidate for military and police work.
What does this mean neurobiologically? German Shepherds often display:
High prey drive: orienting, chasing, gripping behaviors.
Strong defense drive: barking, guarding, territoriality.
Problem-solving intelligence: persistence in tasks requiring mental effort.
Owners who acquire a Shepherd for companionship without providing outlets for these drives often find themselves living with a frustrated, barking, destructive animal
B. Belgian Malinois
The Malinois has been called a “German Shepherd on steroids,” and this is no exaggeration. Selected heavily in military and police programs, the breed was sculpted for extreme intensity, quick arousal, and rapid transitions between aggression and inhibition.
This is a dog whose mesolimbic dopamine system is tuned for perpetual seeking. Malinois thrive when constantly challenged; when deprived of such outlets, they spiral into neurotic behaviors—compulsive spinning, tail chasing, or explosive reactivity.
The tragedy is that Malinois are often marketed as “loyal protectors” or “intelligent companions,” leading unsuspecting owners into disaster.
C. Australian Shepherds and Cattle Dogs
These herding breeds are textbook examples of predatory motor pattern fragmentation. In wolves, the hunting sequence typically unfolds as: orient → eye → stalk → chase → grab-bite → kill-bite → consume. Herding breeds retain the orient, eye, stalk, and chase, but suppress the kill-bite.
Neurobiologically, this means their circuits for orienting and motor control are hyper-efficient. They are compelled to control movement, whether that is livestock, bicycles, children, or cars. Without livestock, they herd shadows, nip heels, or develop obsessive-compulsive behaviors.
D. Siberian Husky
The Husky’s drive is locomotor endurance, encoded in both neural reward pathways and metabolic efficiency. Their brains release dopamine during sustained activity, making running not just enjoyable but neurochemically rewarding.
When confined to small yards or given only brief walks, Huskies become frustrated. They howl, escape, and destroy. This is not disobedience—it is biology rebelling against unmet needs.
E. Catahoula Leopard Dog
Bred for boar hunting and cattle work, Catahoulas embody persistence drive. They were selected to engage prey or livestock despite resistance, sustaining activity in the face of difficulty.
This persistence is underpinned by heritable traits in dopamine receptor density and stress resilience. Owners who underestimate this find themselves with a dog that refuses to quit—whether in chasing prey or chewing through drywall.
Why Drives Cannot Be “Trained Out”
One of the most harmful myths is that training can eliminate drive. Training can shape expression, redirect energy, and create structure. But it cannot remove the underlying motivational circuits.
You cannot train a Husky not to need to run. You can only channel that need through structured outlets.
You cannot train a Malinois not to seek work and challenge constantly. You can only provide controlled scenarios where that drive is expressed productively.
You cannot train a Shepherd not to guard. You can only teach him when guarding is appropriate.
The breeder may have sold you a “companion dog.” The reality is that you have a genetically drivey dog, and the only ethical response is to meet those needs through enrichment, engagement, and respect.
Case Example: The Frustrated Shepherd
Let me illustrate with a real scenario from my work. A family purchased a German Shepherd pup from a breeder who advertised “excellent companion dogs.” By the time the dog was one year old, the family was at their wit’s end. The Shepherd barked incessantly, destroyed furniture, and became reactive on walks.
They arrived at my training center demanding obedience training to “calm him down.” I explained what I am explaining here: you cannot train away genetics. What he needed was a job.
We introduced structured work, tracking exercises, Bite work and advanced obedience integrated with play. Within weeks, the dog’s destructive behaviors diminished—not because the drive disappeared, but because it was finally channeled into tasks aligned with his biology.
This is the core lesson: training cannot erase drive. It can only harness it.
Some trainers claim to “fix” these problems by breaking the dog’s spirit, through harsh corrections, flooding, or relentless suppression. The dog may appear calm, but beneath the surface, what has occurred is not training but trauma.
Suppressing drive at its root is equivalent to inducing learned helplessness, a state where the dog no longer tries because nothing it does matters. Neurobiologically, this corresponds to suppressed dopamine signaling and elevated cortisol. The dog is not calm; it is defeated. As I often tell my clients: the absence of behavior is not proof of training, it is sometimes proof of bad training
So drive is not a nuisance to be trained away. It is the living current of your dog’s genetics and neurobiology. To misunderstand this is to set yourself and your dog up for conflict, frustration, and heartbreak. To respect it is to begin a partnership rooted in truth.
The Neurobiology of Drive and Genetics in Focus
We have explored the ethological foundation of drive, how centuries of selective breeding shaped German Shepherds, Malinois, Huskies, Catahoulas, and herding dogs into the specialists they are today. But to truly grasp why training cannot “erase” drive, we must dive deeper, beneath behavior and even beneath visible temperament, into the neurobiology of drive.
At its root, drive is not an external behavior—it is an internal motivational state regulated by brain circuits, neurotransmitters, and genetic coding. When clients ask me why their Malinois cannot simply relax, or why their Husky is restless even after a two-hour walk, the real answer lies in dopamine receptors, serotonin transporters, and the epigenetic fingerprints left by generations of selection.
Now we will examine:
The dopamine reward system and why high-drive dogs live in constant anticipation.
The roles of serotonin, norepinephrine, and cortisol in modulating drive, stress, and inhibition.
The neuroanatomical circuits linking the limbic brain with cognition, explaining why drivey dogs so easily flip into instinctive states.
The genetic foundations of drive: heritability estimates, candidate genes, and the unintended consequences of selective breeding.
A. Dopamine: The Neurochemical of Drive
Dopamine is often misunderstood as the “pleasure molecule.” In reality, dopamine is the anticipation molecule. It does not flood the brain when the dog achieves a reward but rather when it pursues a reward. This is critical to understanding drive.
In the mesolimbic pathway (ventral tegmental area → nucleus accumbens → prefrontal cortex), dopamine release fuels the sensation of seeking.
High-drive dogs show increased dopaminergic tone—their baseline level of dopamine activity is higher, which keeps them in a perpetual state of readiness
This is why a Belgian Malinois never seems to “turn off.” The act of chasing, biting, searching, or problem-solving is inherently reinforcing—not because of the outcome, but because the pursuit itself triggers dopamine release.
When such a dog is deprived of drive outlets, dopamine dysregulation follows. The result is frustration barking, destructive chewing, compulsive spinning, or explosive outbursts. Owners may perceive this as “misbehavior,” but neurologically it is dopamine seeking an outlet.
B. Serotonin: The Brake on Impulse
Where dopamine propels, serotonin regulates. Originating from the raphe nuclei in the brainstem, serotonin modulates mood, inhibition, and impulse control.
High-drive dogs often exhibit lower baseline serotonergic tone. This is not pathology but adaptation. Lower serotonin makes the animal quicker to act on impulses, less tolerant of inaction, and more prone to explosive responses. In the working arena, this is advantageous—a Malinois that hesitates is a liability.
But in the living room, low serotonergic tone manifests as:
Impatience.
Difficulty settling.
Rapid transitions into arousal states.
Training cannot increase serotonin levels. Only lifestyle management, structured outlets, and, in severe cases, medical intervention can help regulate this neurochemical imbalance.
C. Cortisol and the Stress Axis
Cortisol, produced by the adrenal glands, is the body’s primary stress hormone. Its regulation is controlled by the hypothalamic-pituitary-adrenal (HPA) axis.
When drive is continually suppressed—through lack of outlets, punishment for instinctive behaviors, or chronic frustration—cortisol levels elevate. This has profound consequences:
Hippocampal shrinkage: impairing memory and learning.
Amygdala sensitization: increasing fear and aggression responses.
Immune suppression: reducing health and longevity
This is why dogs raised in environments where their drives are consistently Rewarded often develop anxiety disorders, compulsive behaviors, or aggression.
D. Norepinephrine: The Arousal Regulator
Norepinephrine, closely related to adrenaline, regulates arousal, attention, and readiness for action. In high-drive dogs, norepinephrine systems are primed for hyper-reactivity. This allows a working-line Shepherd to snap into attention at the faintest sound of a threat.
But in the home environment, this same system translates into:
Barking at every noise.
Reactivity on leash.
Inability to relax when stimuli are present.
Again, this is not “bad behavior.” It is the legacy of a norepinephrine system tuned for survival and work.
The Limbic-Cortical Balance: Primal vs. Cognitive Mind
I often explain to my students that dogs operate in two broad states:
The Primal Mind: dominated by the limbic system (amygdala, hypothalamus, basal ganglia). This is where drives, instincts, and emotional reactivity originate.
The Cognitive Mind: involving the prefrontal cortex, hippocampus, and cortical circuits. This is where learning, inhibition, and reflective processing occur.
Drivey dogs, by genetics, spend more time in the Primal Mind. Their neural thresholds for shifting from cognition to instinct are lower. A Malinois may move from engagement to reactivity in fractions of a second, not because it is disobedient, but because its neurobiology prioritizes instinctive action.
Training, therefore, is not about suppressing the Primal Mind but about building bridges into the Cognitive Mind—through engagement, structured outlets, and relationship-based focus.
Genetics in Focus
While neurobiology explains the mechanism, genetics explains the blueprint.
Drive traits are moderately to highly heritable. Studies show:
Herding ability in Border Collies and Aussies: heritability 0.3–0.5.
Protection and defense drive in Shepherds/Malinois: often >0.5.
Endurance drive in Huskies: strongly tied to heritable metabolic efficiency (e.g., mitochondrial adaptations).
This means that even the best training cannot “fix” a mismatch between genetics and lifestyle.
While drive is polygenic, several candidate genes have been consistently implicated:
DRD4 (dopamine receptor D4): Associated with novelty-seeking, attention deficit, and high activity. Variants in DRD4 are common in herding breeds.
SLC6A4 (serotonin transporter): Impacts anxiety, inhibition, and stress resilience. Malinois with certain variants show lower impulse regulation.
MAOA (monoamine oxidase A): Regulates breakdown of serotonin, dopamine, and norepinephrine. Variants here influence aggression and arousal thresholds.
OXTR (oxytocin receptor): Influences bonding, handler focus, and social motivation. Shepherds selected for high handler engagement often carry favorable OXTR variants.
It is not just genes, it is gene expression. Chronic stress, trauma, or enrichment can activate or silence gene expression through epigenetic mechanisms. For example:
Puppies raised without outlets for drive may express higher cortisol reactivity later in life.
Dogs provided with structured drive fulfillment may develop more resilient HPA axes.
This means environment matters—but only within the boundaries set by genetics.
Every breed is the product of human selection pressures that literally sculpted their brains:
Malinois: Selection for military service amplified dopaminergic pursuit and reduced serotonin inhibition, creating intensity and impulsivity.
Huskies: Selection for sledding enhanced mitochondrial efficiency, making endurance not just physical but neurochemically rewarding.
Aussies/Cattle Dogs: Selection fragmented predatory motor patterns, reinforcing orient and chase circuits while suppressing kill-bite.
Catahoulas: Selection for boar hunting amplified persistence and stress resilience genes, ensuring engagement despite resistance.
These are not cosmetic changes. These are neurogenetic rewrites of the animal’s motivational systems.
Owners sometimes ask: “Can’t we just train him to stop being so drivey?” The short answer is no. The long answer is:
You cannot train away dopamine receptor density.
You cannot train away serotonin transporter variants.
You cannot train away a hippocampus sculpted by selection for problem-solving persistence.
What you can do is channel these systems into productive, ethical outlets. But to promise owners that training can “erase” drive is to lie—to them and to the dog
Drive is not an abstract personality quirk. It is the neurobiological and genetic architecture of the dog. From dopamine circuits that thrive on pursuit, to serotonin pathways that regulate inhibition, to the heritable gene variants that sculpt behavior across generations—drive is woven into the dog’s very being.
This truth carries both scientific weight and moral obligation. Owners must understand that when they purchase a high-drive breed, they are inheriting not just a dog, but a neurogenetic blueprint. Trainers must understand that suppression is not training, and that bridging the gap between the Primal and Cognitive mind is the true art of cynology.
Why Suppression Equals Abuse, Applied Training Science, and the Responsibilities of Breeders, Owners, and Trainers
One of the most persistent and dangerous misconceptions in the dog training world is the idea that a drivey dog that lies quietly at your feet is a “calm” dog. Trainers often sell this illusion, and many owners celebrate it as success. But as a cynologist who has devoted decades to the study of canine neuroscience and ethology, I must speak plainly:
When a dog’s genetic drives are suppressed through harsh training methods or chronic deprivation, what occurs is not calmness—it is trauma.
This distinction matters enormously. A dog whose drives are ethically channeled into productive outlets—guided through engagement, meaningful tasks, and fair corrections applied sparingly—becomes calm because its biology has been respected. Its dopamine system is satisfied, its cortisol is regulated, its neuroplasticity remains intact.
By contrast, a dog whose drives have been crushed into silence is “calm” only in the way a prisoner who has surrendered hope is calm. Still on the outside, but broken within.
I will confront the ethical reality of what happens when drives are suppressed, why suppression equals abuse, and where corrections truly fit into the picture—not as weapons to extinguish drives, but as biological feedback tools that must never be the first choice. The truth is clear: drive must first be channeled. Corrections are justified only when a dog, lost in its primal mind, cannot find its way back to cognition without help.
Learned Helplessness: When Spirit Collapses
The collapse induced by harsh suppression is well documented. In the 1960s, psychologist Martin Seligman introduced the concept of learned helplessness through experiments showing that animals exposed to uncontrollable aversive stimuli eventually stopped trying to escape—even when escape became possible.
Translated into dog training, this means that when a dog is punished every time it expresses a genetic behavior—barking, herding, chasing, guarding—it eventually ceases to try at all. The absence of behavior is not a sign of balance. It is a sign of surrender.
Symptoms of Learned Helplessness in Dogs
Lack of initiative: The dog no longer offers behaviors, not even positive ones.
Absence of play: Play drive, fueled by dopamine, evaporates under chronic stress.
Mechanical obedience: Commands are followed, but with no joy or engagement.
Flat affect: The eyes are dull, the tail low, the spark extinguished.
Owners often mistake this state as training success. But what they are really witnessing is the collapse of exploratory behavior, the extinguishing of drive, the surrender of will.
This is not training. This is breaking.
Neuroendocrine Damage of Suppression
The damage inflicted by suppression is not only psychological but neurological. Chronic suppression rewires the brain and body through the neuroendocrine system, with long-lasting consequences.
A. Elevated Cortisol
Suppression chronically activates the hypothalamic-pituitary-adrenal (HPA) axis, flooding the body with cortisol. Over time, elevated cortisol produces:
Hippocampal shrinkage, impairing learning and spatial memory.
Amygdala hyperactivity, heightening fear and aggression responses.
Immune suppression, reducing resilience and increasing illness risk.
B. Suppressed Dopamine
When drive is consistently thwarted, dopamine release diminishes. Without dopamine:
Motivation collapses.
Neuroplasticity declines.
Joy and initiative fade.
C. Reduced Neuroplasticity
Neuroplasticity—the brain’s ability to adapt through experience—depends on balanced dopamine and stress regulation. Chronic suppression locks the dog into rigid patterns of fear or apathy.
Thus, a dog whose drives are crushed is not just emotionally harmed. It is neurologically altered.
The Illusion of Calmness
A suppressed dog may indeed appear calm. It may lie quietly, make no attempt to play, and show no outward distress. To the untrained eye, this looks like obedience and balance. Neuroscience reveals otherwise.
This is not calmness born of satisfaction. It is the stillness of defeat.
Contrast with True Calmness
Suppressed calmness is absence: no play, no initiative, no drive.
True calmness is presence: joy, engagement, controlled energy.
A genuine working partnership is not defined by silence. It is defined by energy channeled into purpose, not energy beaten into submission.
Corrections: Biology, Not Cruelty
One of the most polarizing debates in dog training is over corrections. Many equate correction with cruelty. Others wield correction as their primary tool. Both extremes miss the truth. Harsh suppression that seeks to extinguish drive is unquestionably abuse. But fair, well-timed corrections are not only ethical—they are biological.
Across the animal kingdom, corrections are how boundaries are established and learning occurs.
Wolf Packs: Pups are corrected for biting too hard, for trespassing social boundaries, for failing to heed signals. These corrections are sharp, immediate, and proportionate—but they are not designed to break the pup’s spirit. They teach the pup how to function in the group.
Mother Dogs: A dam will growl, snap, or physically block a puppy that becomes too rough. The correction is swift, clear, and then immediately over. The pup learns through this feedback loop what behaviors are acceptable.
Wild Primates: Grooming interruptions, vocal reprimands, or even physical shoves are part of normal social corrections in primates.
Humans: Children are corrected by parents and peers. Not all correction is punishment; it is a form of feedback that shapes behavior and social integration.
In other words: correction is woven into the biological fabric of social species. To deny its place is to deny dogs a form of communication they innately understand. The problem is not correction itself—it is when correction is misused to crush drives rather than guide them.
The Neurobiology of Correction
From a neuroscience perspective, fair corrections act not as punishments but as interruptors.
A mild aversive stimulus (verbal interrupter, leash guidance, spatial pressure) briefly engages the amygdala.
This arousal creates a pause in limbic-driven behavior.
If immediately followed by redirection into engagement or task, the prefrontal cortex activates.
The dog makes a cognitive choice: revert to instinct or re-engage with the handler.
High-drive dogs often spend more time in their primal mind (amygdala-driven, instinctive, reactive). Corrections are biological bridges that pull them momentarily out of that limbic loop, opening a window for cognition.
The Yerkes-Dodson Law (1908) and later work by Arnsten (2009) show that moderate stress enhances prefrontal activation and learning, while overwhelming stress collapses cognition. Mild, fair corrections fall into the former category. Harsh suppression falls into the latter, inducing trauma.
Thus, ethical corrections enhance learning. Harsh suppression destroys it.
Why Corrections Should Never Be First Choice
Drive Must Be Channeled First
The first responsibility is always to channel drive through enrichment, outlets, and engagement.
Dopamine is released during pursuit, not just reward (Berridge & Robinson, 1998).
Enrichment lowers cortisol and builds resilience (Hennessy et al., 1998, shelter dog studies).
Without outlets, corrections are futile. They may stop behavior in the moment, but the underlying neurochemical imbalance—dopamine seeking and cortisol rising—remains unresolved.
Corrections should never be the foundation. They are the safety rails, not the road itself. Used ethically, they interrupt primal overdrive, but the true solution is always found in drive fulfillment and engagement.
Applied Training Science: The Ethical Alternative
If suppression is abuse, the ethical alternative is applied neuroscience-based training, where the goal is not to erase drive but to redirect it into engagement and meaningful outlets.
Engagement makes the handler the center of meaning. It is not bribery. It is the neurochemical linking of the dog’s dopamine system to the handler’s presence and actions. Through consistent reinforcement and oxytocin-mediated bonding, the dog learns: “My drives are best fulfilled with you.”
Many high-drive breeds are cognitively gifted. Without intellectual outlets, they spiral into neuroticism. Tasks such as:
Scent work.
Tracking and search games.
Puzzle feeders and shaping exercises.
By engaging the prefrontal cortex, these tasks balance primal and cognitive states.
Physical Outlets
Drive expression must align with genetics:
Huskies: structured running, canicross, bikejoring.
Malinois: bite work, tactical obedience.
Shepherds: tracking, protection, advanced obedience.
Aussies/Cattle Dogs: agility, herding games.
Catahoulas: endurance hiking, scent trailing.
The point is not exhaustion but fulfillment.
Neuroscience shows that predictability stabilizes the HPA axis. Clear boundaries and routines reduce stress, while chaos amplifies it. Drivey dogs need structure as much as outlets.
Responsibilities Across the Chain
A. Breeders
Much frustration begins here. Too many breeders misrepresent high-drive dogs as simple companions. Ethical breeders must:
Be transparent about drive levels.
Place dogs with suitable owners.
Avoid exaggerating drive to extremes for image or sport.
To advertise a Malinois as a cuddly lapdog is not just dishonest. It is dangerous.
B. Owners
Owning a drivey dog is a lifestyle. Owners must accept:
Drive does not “settle with age.”
Obedience alone is insufficient.
The responsibility is lifelong.
When owners honor this, frustration becomes partnership. When they deny it, the dog suffers.
C. Trainers
Too many trainers sell “calmness packages” that amount to suppression. This is malpractice. Ethical trainers must:
Educate clients about biology.
Refuse suppression-based solutions.
Model respect for genetics and drives.
To promise to erase a Malinois’s drive is not training. It is complicity in abuse.
Conclusion:
The refrain I want every owner of a drivey dog to remember is this:
Your dog is not broken. Your expectations are.
You cannot ask a Husky not to run. You cannot ask a Malinois not to work. You cannot ask a Shepherd not to guard. These are not quirks. They are the living expression of genes sculpted by centuries of selection.
The art of training is not to erase these drives but to harness them into partnership.
When we respect the truth of genetics, when we honor the neurobiology of drive, we unlock harmony—not through suppression, but through understanding.
This is the essence of my work, and the legacy I aim to leave: to shift the conversation from control to respect, from breaking to guiding, from ignorance to science.
The dog does not exist to fulfill human fantasies of convenience. The dog exists as a being of biology, drive, and truth. To love the dog is to accept this truth and rise to meet it.
Bart de Gols
References:
Learned Helplessness & Stress
Seligman, M. E. P. (1972). Learned helplessness. Annual Review of Medicine, 23(1), 407–412.
Maier, S. F., & Seligman, M. E. (2016). Learned helplessness at fifty: Insights from neuroscience. Psychological Review, 123(4), 349–367.
Hennessy, M. B., Williams, M. T., Miller, D. D., Douglas, C. W., & Voith, V. L. (1998). Influence of male and female petters on plasma cortisol and behaviour: Can human interaction reduce the stress of dogs in a public animal shelter? Applied Animal Behaviour Science, 61(1), 63–77.
Neuroendocrine Responses: Cortisol, Dopamine, Neuroplasticity
Arnsten, A. F. T. (2009). Stress signalling pathways that impair prefrontal cortex structure and function. Nature Reviews Neuroscience, 10(6), 410–422.
McEwen, B. S. (2007). Physiology and neurobiology of stress and adaptation: Central role of the brain. Physiological Reviews, 87(3), 873–904.
Sapolsky, R. M. (2004). Why zebras don’t get ulcers: The acclaimed guide to stress, stress-related diseases, and coping. Holt Paperbacks.
Dias-Ferreira, E., Sousa, J. C., Melo, I., et al. (2009). Chronic stress causes frontostriatal reorganization and affects decision-making. Science, 325(5940), 621–625.
Dopamine, Motivation & Drive
Berridge, K. C., & Robinson, T. E. (1998). What is the role of dopamine in reward: Hedonic impact, reward learning, or incentive salience? Brain Research Reviews, 28(3), 309–369.
Schultz, W. (2007). Multiple dopamine functions at different time courses. Annual Review of Neuroscience, 30, 259–288.
Salamone, J. D., & Correa, M. (2012). The mysterious motivational functions of mesolimbic dopamine. Neuron, 76(3), 470–485.
Serotonin & Impulse Regulation
Seo, D., Patrick, C. J., & Kennealy, P. J. (2008). Role of serotonin and dopamine system interactions in the neurobiology of impulsive aggression and its comorbidity with other clinical disorders. Aggression and Violent Behavior, 13(5), 383–395.
Lesch, K. P., Bengel, D., Heils, A., et al. (1996). Association of anxiety-related traits with a polymorphism in the serotonin transporter gene regulatory region. Science, 274(5292), 1527–1531.
Norepinephrine, Arousal, and Attention
Aston-Jones, G., & Cohen, J. D. (2005). An integrative theory of locus coeruleus–norepinephrine function: Adaptive gain and optimal performance. Annual Review of Neuroscience, 28, 403–450.
Yerkes–Dodson Law & Stress-Learning Relationship
Yerkes, R. M., & Dodson, J. D. (1908). The relation of strength of stimulus to rapidity of habit‐formation. Journal of Comparative Neurology and Psychology, 18(5), 459–482.
Arnsten, A. F. T. (2009). (listed above).
Canine Genetics & Heritability of Behavior
Overall, K. L. (2013). Manual of clinical behavioral medicine for dogs and cats. Elsevier.
Ilska, J., Haskell, M. J., Blott, S. C., Sánchez-Molano, E., Polgar, Z., Lofgren, S. E., … & Wiener, P. (2017). Genetic characterization of dog personality traits. Genetics, 206(2), 1101–1111.
Chase, K., et al. (2009). Genetic basis for systems of canine behavior: I. heritability of fear and aggression in German Shepherd Dogs. Applied Animal Behaviour Science, 120(1–2), 125–135.
Persson, M. E., Roth, L. S., Johnsson, M., Wright, D., & Jensen, P. (2015). Human-directed social behaviour in dogs shows significant heritability. Genes, Brain and Behavior, 14(4), 337–344.
Oxytocin & Engagement with Handlers
Nagasawa, M., Kikusui, T., Onaka, T., & Ohta, M. (2009). Dog’s gaze at its owner increases owner’s urinary oxytocin during social interaction. Hormones and Behavior, 55(3), 434–441.
Kis, A., Bence, M., Lakatos, G., Pergel, E., Turcsán, B., Pluijmakers, J., … & Kubinyi, E. (2014). Oxytocin receptor gene polymorphisms are associated with human directed social behavior in dogs (Canis familiaris). PLoS One, 9(1), e83993.
Applied Dog Training & Welfare Studies
Hiby, E. F., Rooney, N. J., & Bradshaw, J. W. (2004). Dog training methods: Their use, effectiveness and interaction with behaviour and welfare. Animal Welfare, 13(1), 63–70.
Rooney, N. J., & Cowan, S. (2011). Training methods and owner–dog interactions: Links with dog behaviour and learning ability. Applied Animal Behaviour Science, 132(3–4), 169–177.
Blackwell, E. J., Twells, C., Seawright, A., & Casey, R. A. (2008). The relationship between training methods and the occurrence of behavior problems, as reported by owners, in a population of domestic dogs. Journal of Veterinary Behavior, 3(5), 207–217.
Ethology, Predatory Motor Patterns, Breed Selection
Coppinger, R., & Coppinger, L. (2001). Dogs: A new understanding of canine origin, behavior, and evolution. University of Chicago Press.
Mech, L. D., & Boitani, L. (2003). Wolves: Behavior, ecology, and conservation. University of Chicago Press.
Goodwin, D., Bradshaw, J. W. S., & Wickens, S. M. (1997). Paedomorphosis affects agonistic visual signals of domestic dogs. Animal Behaviour, 53(2), 297–304.