Dog behavior

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Dogs roughhousing
A drawing by Konrad Lorenz showing facial expressions of a dog - a communication behavior

Dog behavior has been shaped by millennia of contact with humans. As a result of this physical and social evolution, dogs, more than any other species, have acquired the ability to understand and communicate with humans and they are uniquely attuned to our behaviors.[1] Behavioral scientists have uncovered a wide range of social-cognitive abilities in the domestic dog. Dog behavior is the internally coordinated responses of individuals or groups of domestic dogs to internal and external stimuli.[2]

Co-evolution with humans[edit]

The origin of the domestic dog (Canis lupus familiaris or Canis familiaris) is not clear. Whole genome sequencing indicates that the dog, the gray wolf and the extinct Taymyr wolf diverged at around the same time 27,000-40,000 years ago.[3] How dogs became domesticated is not clear, however the two main hypotheses are self-domestication or human domestication. There exists evidence of human-canine coevolution.


Further information: Dog intelligence



Further information: Dog anatomy § Senses

The dog's senses include vision, hearing, sense of smell, taste, touch and sensitivity to the earth's magnetic field.

Communication behavior[edit]

Further information: Dog communication


Social behavior[edit]



Dog playing with a guinea pig.

Play between dogs usually involves several behaviours that are often seen in aggressive encounters, for example, nipping, biting and growling. It is therefore important for the dogs to place these behaviours in the context of play, rather than aggression. Dogs signal their intent to play with a range of behaviours including a "play-bow", "face-paw" "open-mouthed play face" and postures inviting the other dog to chase the initiator. Similar signals are given throughout the play bout to maintain the context of the potentially aggressive activities.[5]




It is a common misconception that winning and losing games such as "tug-of-war" and "rough-and-tumble" can influence a dog's dominance relationship with humans. Rather, the way in which dogs play indicates their temperament and relationship with their owner. Dogs that play rough-and-tumble are more amenable and show lower separation anxiety than dogs which play other types of games, and dogs playing tug-of-war and "fetch" are more confident. Dogs which start the majority of games are less amenable and more likely to be aggressive.[8]

Playing with humans can affect the cortisol levels of dogs. In one study, the cortisol responses of police dogs and border guard dogs was assessed after playing with their handlers. The cortisol concentrations of the police dogs increased, whereas the border guard dogs' hormone levels decreased. The researchers noted that during the play sessions, police officers were disciplining their dogs, whereas the border guards were truly playing with them, i.e. this included bonding and affectionate behaviours. They commented that several studies have shown that behaviours associated with control, authority or aggression increase cortisol, whereas play and affiliative behaviour decrease cortisol levels.[9]



A study found a third of dogs suffered from anxiety when separated from others.[11]


Two Labrador Dog Fighting

The term personality has been applied to human research, whereas the term temperament has been mostly used for animal research.[12] However, both terms have been used interchangeably in the literature, or purely to distinguished humans for animals and avoid anthropomorphism.[13] Personality can be defined by “a set of behaviours that are consistent over context and time”.[14] Human personality is often studied using models that look at broad dimensions of personality. For example, the Five Factor Model is one of the most commonly used model, and the most extensively studied. It is composed of five dimensions: openness, conscientiousness, extraversion, agreeableness, and neuroticism.[15] Studies of dog personality have also tried to identify the presence of broad personality traits that are stable over time.[13][14] Recently, dog’s personality dimension has been shown to be relatively consistent over time.[16][17]

There are different approaches to assess dog personality:

  • Ratings of individual dogs: either a caretaker or a dog expert who is familiar with the dog is asked to answer a questionnaire, for instance the Canine Behavioural Assessment and Research Questionnaire,[18] concerning how often the dog show certain type of behaviour.
  • Test batteries: the dog is submitted to a set of tests and its reactions are evaluated on a behavioural scale. For instance, the dog is presented to a familiar and then an unfamiliar person :in order to measure sociability or aggression.[19]
  • Observational test: The dog’s behaviour is evaluated in a selected but not controlled environment. An observer focus on the dog’s reactions to naturally occurring stimuli. For example a :walk through the supermarket can allow the observer to see the dog in various types of conditions (crowd, loud noise …).[20]

Several potential personality traits have been identified in dogs, for instance "Playfulness", "Curiosity/Fearlessness, "Chase-proneness", "Sociability and Aggressiveness" and "Shyness–Boldness".[21][22] A meta-analysis of 51 published peer reviewed articles identified seven dimension of canine personality:[13]

  1. Reactivity (approach or avoidance of new objects, increased activity in novel situations)
  2. Fearfulness (shaking, avoiding novel situations)
  3. Activity
  4. Sociability (initiating friendly interactions with people and other dogs)
  5. Responsiveness to Training (working with people, learning quickly)
  6. Submissiveness
  7. Aggression

Dog Breed plays an important role in the dog’s personality dimensions,[23][24] while the effects of age and sex have not been clearly determined.[14] Dogs personality models can be used for a range of tasks, including guide and working dog selection, finding appropriate families to re-home shelter dogs, or selecting breeding stock.[25][26][27]

Leadership, dominance and social groups[edit]

Two dogs playing follow the leader.

Dominance is a descriptive term for the relationships between pairs of individuals. Among ethologists, dominance has been defined as ‘‘an attribute of the pattern of repeated, agonistic interactions between two individuals, characterized by a consistent outcome in favor of the same dyad member and a default yielding response of its opponent rather than escalation. The status of the consistent winner is dominant and that of the loser subordinate.’’[28] Another definition is that a dominant animal has priority of access to resources.[28] Dominance is a relative attribute, not absolute; there is no reason to assume that a high-ranking individual in one group would also become high ranking if moved to another. Nor is there any good evidence that ‘‘dominance’’ is a lifelong character trait. Competitive behavior characterized by confident (e.g. growl, inhibited bite, stand over, stare at, chase, bark at) and submissive (e.g. crouch, avoid, displacement lick/yawn, run away) patterns exchanged.[29]

One test to ascertain which in a group was the dominant dog used the following criteria: When a stranger comes to the house, which dog starts to bark first or if they start to bark together, which dog barks more or longer? Which dog licks more often the other dog’s mouth? If the dogs get food at the same time and at the same spot, which dog starts to eat first or eats the other dog’s food? If the dogs start to fight, which dog wins usually?[30]


Two dogs that are contesting possession of a highly valued resource for the first time, if one is in a state of emotional arousal, or if one is in pain, or if reactivity is influenced by recent endocrine changes or motivational states such as hunger, then the outcome of the interaction may be different than if none of these factors were present. Equally, the threshold at which aggression is shown may be influenced by a range of medical factors, or, in some cases, precipitated entirely by pathological disorders. Hence, the contextual and physiological factors present when 2 dogs first encounter each other may profoundly influence the long-term nature of the relationship between those dogs. The complexity of the factors involved in this type of learning means that dogs may develop different ‘‘expectations’’ about the likely response of another individual for each resource in a range of different situations. Puppies learn early not to challenge an older dog and this respect stays with them into adulthood. When adult animals meet for the first time, they have no expectations of the behavior of the other: they will both, therefore, be initially anxious and vigilant in this encounter (characterized by the tense body posture and sudden movements typically seen when 2 dogs first meet), until they start to be able to predict the responses of the other individual. The outcome of these early adult–adult interactions will be influenced by the specific factors present at the time of the initial encounters. As well as contextual and physiological factors, the previous experiences of each member of the dyad of other dogs will also influence their behavior.[29]




Further information: Dog communication

Feral dogs[edit]



See further: behavior compared to other canids.

Other behavior[edit]

Dogs have a general behavioral trait of strongly preferring novelty ("neophillia") compared to familiarity.[34] The average sleep time of a dog in captivity in a 24-hour period is 10.1 hours.[35]

Reproduction behavior[edit]

Further information: Canine reproduction

Estrous cycle and mating[edit]

Although puppies do not have the urge to procreate, males sometimes engage in sexual play in the form of mounting, as early as 5 weeks-of-age.

Dogs reach sexual maturity and can reproduce during their first year, in contrast to wolves at two years-of-age. Bitches have their first estrus ("heat") at 6 to 12 months-of-age; smaller dogs tend to come into heat earlier whereas larger dogs take longer to mature.

Dog bitches have an estrous cycle that is nonseasonal and monestrus, i.e. there is only one estrus per estrous cycle. The interval between one estrus and another is, on average, seven months, however, this may range between 4 and 12 months. This interestrous period is not influenced by the photoperiod or pregnancy. The average duration of estrus is 9 days with spontaneous ovulation usually about 3 days after the onset of estrus.[36]

For several days before estrus, a phase called proestrus, the bitch may show greater interest in male dogs and "flirt" with them (proceptive behavior). There is progressive vulval swelling and some bleeding. If males try to mount a bitch during proestrus, she may avoid mating by sitting down or turning round and growling or snapping.

Estrous behavior in the bitch is usually indicated by her standing still with the tail held up, or to the side of the perineum, when the male sniffs the vulva and attempts to mount. This tail position is sometimes called “flagging”. The bitch may also turn, presenting the vulva to the male.[36]

The male dog mounts the female and is able to achieve intromission with a non-erect penis, which contains a bone called the os penis. The dog's penis enlarges inside the vagina, thereby preventing its withdrawal; this is sometimes known as the “tie” or “copulatory lock”. The male dog rapidly thrust into the female for 1–2 minutes then dismounts with the erect penis still inside the vagina, and turns to stand rear-end to rear-end with the bitch for up to 30 to 40 minutes; the penis is twisted 180 degrees in a lateral plane. During this time, prostatic fluid is ejaculated.[36]


Parenting and early life[edit]


Behavior problems[edit]


Separation anxiety[edit]

When dogs are separated from humans, usually the owner, they often display behaviours such as destructiveness, faecal or urinary elimination, hypersalivation or vocalisation. Dogs from single-owner homes are approximately 2.5 times more likely to have separation anxiety compared to dogs from multiple-owner homes. Furthermore, sexually intact dogs are only one third as likely to have separation anxiety as neutered dogs. The sex of dogs and whether there is another pet in the home do not have an effect on separation anxiety.[39]

Noise anxiety[edit]


Tail chasing[edit]

Tail chasing can be classified as a stereotypy. In one clinical study on this potential behavioral problem, 18 tail-chasing terriers were given clomipramine orally at a dosage of 1 to 2 mg/kg (0.5 to 0.9 mg/lb) of body weight, every 12 hours. Three of the dogs required treatment at a slightly higher dosage range to control tail chasing, however, after 1 to 12 weeks of treatment, 9 of 12 dogs were reported to have a 75% or greater reduction in tail chasing.[40]

Behavior compared to other canids[edit]

Comparisons made within the wolf-like canids allow the identification of those behaviors that may have been inherited from common ancestry and those that may have been the result of domestication or other relatively recent environmental changes.[32] Studies of free-ranging African Basenjis and New Guinea Singing Dogs indicate that their behavioral and ecological traits were the result of environmental selection pressures or selective breeding choices and not the result of artificial selection imposed by humans.[41]


Unlike other domestic species which were primarily selected for production-related traits, dogs were initially selected for their behaviors.[42][43] In 2016, a study found that there were only 11 fixed genes that showed variation between wolves and dogs. These gene variations were unlikely to have been the result of natural evolution, and indicate selection on both morphology and behavior during dog domestication. These genes have been shown to affect the catecholamine synthesis pathway, with the majority of the genes affecting the fight-or-flight response[43][44] (i.e. selection for tameness), and emotional processing.[43] Dogs generally show reduced fear and aggression compared to wolves.[43][45] Some of these genes have been associated with aggression in some dog breeds, indicating their importance in both the initial domestication and then later in breed formation.[43]

Social structure[edit]

Among canids, packs are the social units that hunt, rear young and protect a communal territory as a stable group and their members are usually related.[46] Members of the feral dog group are usually not related. Feral dog groups are composed of a stable 2-6 members compared to the 2-15 member wolf pack whose size fluctuates with the availability of prey and reaches a maximum in winter time. The feral dog group consists of monogamous breeding pairs compared to the one breeding pair of the wolf pack. Agonistic behavior does not extend to the individual level and does not support a higher social structure compared to the ritualized agonistic behavior of the wolf pack that upholds its social structure. Feral pups have a very high mortality rate that adds little to the group size, with studies showing that adults are usually killed through accidents with humans, therefore other dogs need to be co-opted from villages to maintain stable group size.[32]


The critical period for socialization begins with walking and exploring the environment. Dog and wolf pups both develop the ability to see, hear and smell at 4 weeks of age. Dogs begin to explore the world around them at 4 weeks of age with these senses available to them, while wolves begin to explore at 2 weeks of age when they have the sense of smell but are functionally blind and deaf. The consequences of this is that more things are novel and frightening to wolf pups. The critical period for socialization closes with the avoidance of novelty, when the animal runs away from - rather than approaching and exploring - novel objects. For dogs this develops between 4 and 8 weeks of age. Wolves reach the end of the critical period after 6 weeks, after which it is not possible to socialize a wolf.[37]

Dog puppies require as little as 90 minutes of contact with humans during their critical period of socialization to form a social attachment. This will not create a highly social pet but a dog that will solicit human attention.[47] Wolves require 24 hours contact a day starting before 3 weeks of age. To create a socialized wolf the pups are removed from the den at 10 days of age, kept in constant human contact until they are 4 weeks old when they begin to bite their sleeping human companions, then spend only their waking hours in the presence of humans. This socialization process continues until age 4 months, when the pups can join other captive wolves but will require daily human contact to remain socialized. Despite this intensive socialization process, a well-socialized wolf will behave differently to a well-socialized dog and will display species-typical hunting and reproductive behaviors, only closer to humans than a wild wolf. These wolves do not generalize their socialization to all humans in the same manner as a socialized dog and they remain more fearful of novelty compared to socialized dogs.[48]



Despite claims that dogs show more human-like social cognition than wolves,[50][51][52] several recent studies have demonstrated that if wolves are properly socialized to humans and have the opportunity to interact with humans regularly, then they too can succeed on some human-guided cognitive tasks,[53][54][55][56][57] in some cases out-performing dogs at an individual level.[58] Similar to dogs, wolves can also follow more complex point types made with body parts other than the human arm and hand (e.g. elbow, knee, foot).[57] Both dogs and wolves have the cognitive capacity for prosocial behavior toward humans; however it is not guaranteed. For canids to perform well on traditional human-guided tasks (e.g. following the human point) both relevant lifetime experiences with humans - including socialization to humans during the critical period for social development - and opportunities to associate human body parts with certain outcomes (such as food being provided by human hands, a human throwing or kicking a ball, etc.) are required.[59]



Dogs reach sexual maturity and can reproduce during their first year in contrast to a wolf at two years. The female dog can bear another litter within 8 months of the last one. The canid genus is influenced by the photoperiod and generally reproduces in the springtime.[32] Domestic dogs are not reliant on seasonality for reproduction in contrast to the wolf, coyote, Australian dingo and African basenji that may have only one, seasonal, estrus each year.[36] Feral dogs are influenced by the photoperiod with around half of the breeding females mating in the springtime, which is thought to indicate an ancestral reproductive trait not overcome by domestication,[32] as can be inferred from wolves[60] and Cape hunting dogs.[61]

Domestic dogs are polygamous in contrast to wolves that are generally monogamous. Therefore, domestic dogs have no pair bonding and the protection of a single mate, but rather have multiple mates in a year. There is no paternal care in dogs as opposed to wolves where all pack members assist the mother with the pups. The consequence is that wolves put a lot of energy into producing a few pups in contrast to dogs that maximize the production of pups. This higher pup production rate enables dogs to maintain or even increase their population with a lower pup survival rate than wolves, and allows dogs a greater capacity than wolves to grow their population after a population crash or when entering a new habitat. It is proposed that these differences are an alternative breeding strategy adapted to a life of scavenging instead of hunting.[37] In contrast to domestic dogs, feral dogs are monogamous. Domestic dogs tend to have a litter size of 10, wolves 3, and feral dogs 5-8. Feral pups have a very high mortality rate with only 5% surviving at the age of one year, and sometimes the pups are left unattended making them vulnerable to predators.[32] Domestic dogs stand alone among all canids for a total lack of paternal care.[62]

Dogs differ from wolves and most other large canid species as they generally do not regurgitate food for their young, nor the young of other dogs in the same territory.[63] However, this difference was not observed in all domestic dogs. Regurgitating of food by the females for the young, as well as care for the young by the males, has been observed in domestic dogs, dingos and in feral or semi-feral dogs. In one study of a group of free-ranging dogs, for the first 2 weeks immediately after parturition the lactating females were observed to be more aggressive to protect the pups. The male parents were in contact with the litters as ‘guard’ dogs for the first 6–8 weeks of the litters’ life. In absence of the mothers, they were observed to prevent the approach of strangers by vocalizations or even by physical attacks. Moreover, one male fed the litter by regurgitation showing the existence of paternal care in some free-roaming dogs[64]




Despite claims in the popular press, studies could not find evidence of a single predation on cattle by feral dogs.[32][65][66] However, domestic dogs were responsible for the death of 3 calves over one 5-year study.[66] Other studies in Europe and North America indicate only limited success in the consumption of wild boar, deer and other ungulates, however it could not be determined if this was predation or scavenging on carcasses. Studies have observed feral dogs conducting brief, uncoordinated chases of small game with constant barking - a technique without success.[32]

In 2004, a study reviewed 5 other studies of feral dogs published between 1975 and 1995 and concluded that their pack structure is very loose and rarely involves any cooperative behavior, either in raising young or in obtaining food.[67] Feral dogs are primarily scavengers, with studies showing that unlike their wild cousins, they are poor ungulate hunters, having little effect on wildlife populations where they are sympatric.[68]:267 However, several garbage dumps located within the feral dog's home range are important for their survival.[69]

Dogs in human society[edit]

Further information: Human-canine bond

Dogs likely were the first animals to be domesticated and as such have shared a common environment with humans for over ten thousand years. Only recently, however, has this species' behavior been subject to scientific scrutiny. Most of this work has been inspired by research in human cognitive psychology and suggests that in many ways dogs are more human-like than any other species, including nonhuman primates.[70][copyright violation?]

Studies using an operant framework have indicated that humans can influence the behavior of dogs through food, petting and voice. Food and 20–30 seconds of petting maintained operant responding in dogs.[71] Some dogs will show a preference for petting once food is readily available, and dogs will remain in close proximity to a person providing petting and show no satiation to that stimulus.[72] Petting alone was sufficient to maintain the operant response of military dogs to voice commands, and responses to basic obedience commands in all dogs increased when only vocal praise was provided for correct responses.[73]



Dogs in the family[edit]

Further information: Companion dog

Dogs at work[edit]

Service dogs are those that are trained to help people with disabilities. Detection dogs are trained to using their sense of smell to detect substances such as explosives, illegal drugs, wildlife scat, or blood. In science, dogs have helped humans understand about the conditioned reflex. Attack dogs, dogs that have been trained to attack on command, are employed in security, police, and military roles.


Main article: Dog attack
A dog's teeth can inflict serious injuries

The human-dog relationship is based on unconditional trust, however if this trust is lost it will be difficult to reinstate. As a last resort, humans will use a slap but a dog will use a bite. A dog's thick fur protects it from the bite of another dog but humans are furless and are not so protected.[77]

In the UK between 2005 and 2013, there were 17 fatal dog attacks. In 2007-08 there were 4,611 hospital admissions due to dog attacks, which increased to 5,221 in 2008-09. It has been estimated that more than 200,000 people a year are bitten by dogs in England, with the annual cost to the National Health Service of treating injuries about £3 million.[78] A report published in 2014 stated there were 6,743 hospital admissions specifically caused by dog bites, a 5.8% increase from the 6,372 admissions in the previous 12 months.[79]

In the US between 1979 and 1996, there were more than 300 human dog bite-related fatalities.[80] In the US in 2013, there were 31 dog-bite related deaths. Each year, more than 4.5 million people in the US are bitten by dogs and almost 1 in 5 require medical attention.[81]

Attack training is condemned by some as promoting ferocity in dogs; a 1975 American study showed that 10% of dogs that have bitten a person received attack dog training at some point.[82]

See also[edit]


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Further reading[edit]

  • Hare, Brian & Woods, Venessa. The Genius of Dogs (2013 Penguin Publishing Group) Reveals research findings about how dogs think and how we humans can have deeper relationships with them.
  • Miklosi, Adam. Dog Behaviour, Evolution, and Cognition (2007 Oxford University Press) Provides a basis for a complete dog behavioural biology based on concepts derived from contemporary ethology.
  • Pet Behavior articles from the ASPCA

External links[edit]