Dog behavior

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

Dog intelligence is the ability of the dog to perceive information and retain it as knowledge in order to solve problems. Dogs have been shown to learn by inference. A study with Rico showed that he knew the labels of over 200 different items. He inferred the names of novel items by exclusion learning and correctly retrieved those novel items immediately. He also retained this ability four weeks after the initial exposure. Dogs have advanced memory skills. A study documented the learning and memory capabilities of a border collie, "Chaser", who had learned the names and could associate by verbal command over 1,000 words. Dogs are able to read and react appropriately to human body language such as gesturing and pointing, and to understand human voice commands. After undergoing training to solve a simple manipulation task, dogs that are faced with an insolvable version of the same problem look at the human, while socialized wolves do not. Dogs demonstrate a theory of mind by engaging in deception. ==Senses==

The dog's senses include vision, hearing, sense of smell, taste, touch, proprioception, and sensitivity to the Earth's magnetic field. ==Communication behavior==

Dog communication is about how dogs "speak" to each other, how they understand messages that humans send to them, and how humans can translate the ideas that dogs are trying to transmit. ==Social behavior==

Two studies have indicated that dog behavior vary based on their size, body weight, and skull size. Play Dog and dog and maltipoo running back while playing fetch Play between dogs usually involves several behaviors often seen in aggressive encounters, such as nipping, biting and growling. It is therefore important for the dogs to place these behaviors in the context of the play, rather than aggression. Dogs signal their intent to play with a range of behaviors 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 to maintain the context of the potentially aggressive activities. From a young age, dogs engage in play with one another. Dog play is made up primarily of mock fights. It is believed that this behavior, which is most common in puppies, is training for important behaviors later in life. Some early socialization methods, such as puppy classes or "puppy parties" have been shown to positively influence adult dog behavior. Play between puppies is not necessarily a 50:50 symmetry of dominant and submissive roles between the individuals; dogs who engage in greater rates of dominant behaviors (e.g. chasing, forcing partners down) at later ages also initiate play at higher rates. This could imply that winning during play becomes more important as puppies mature. Emotional contagion is linked to facial mimicry in humans and primates. Facial mimicry is an automatic response that occurs in less than 1 second in which one person involuntarily mimics another person's facial expressions, forming empathy. It has also been found in dogs at play, and play sessions lasted longer when there were facial mimicry signals from one dog to another. Dog and human with his dogs Jake and Scout The motivation for a dog to play with another dog is different from that of a dog playing with a human. Dogs walked together with opportunities to play with one another and play with their owners, choose to play with their owners at the same frequency as dogs being walked alone. Dogs in households with two or more dogs only play slightly more often with their owners than dogs in households with a single dog, indicating the motivation to play with other dogs does not substitute for the motivation to play with humans. It is a common misconception that winning and losing games such as "tug-of-war" and "rough-and-tumble" can influence a dog's dominant relationship with humans. Rather, how 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 that start most games are less amenable and more likely to be aggressive. Scientific studies highlight the unique bonds between humans and dogs, showing how friendship and communication has evolved through domestication. Playing with humans can affect the cortisol levels of dogs. In one study, the cortisol responses of police dogs and border guard dogs were 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 behaviors. They commented that several studies have shown that behaviors associated with control, authority or aggression increase cortisol, whereas play and affiliation behavior decrease cortisol levels. Empathy In 2012, a study found that dogs oriented toward their owner or a stranger more often when the person was pretending to cry than when they were talking or humming. When the stranger pretended to cry, rather than approaching their usual source of comfort, their owner, dogs sniffed, nuzzled and licked the stranger instead. The dogs' pattern of response was behaviorally consistent with an expression of empathic concern. Personalities The term personality has been applied to human research, whereas the term temperament has been mostly used for animal research. A meta-analysis of 51 published peer-reviewed articles identified seven dimensions of canine personality: However, a study in December 2022 challenged those findings after the researching of the genetic codes of 4,000 dogs and 46,000 dog owners and concluded that a dog's breed does genetically influence a dog's personality. The effects of age and sex have not been clearly determined. The anal gland of canines give a particular signature to fecal deposits and identifies the marker as well as the place where the dung is left. Dogs are very particular about these landmarks, and engage in what is to humans a meaningless and complex ritual before defecating. Most dogs start with a careful bout of sniffing of a location, perhaps to erect an exact line or boundary between their territory and another dog's territory. This behavior may also involve a small degree of elevation, such as a rock or fallen branch, to aid scent dispersal. Scratching the ground after defecating is a visual sign pointing to the scent marking. The freshness of the scent gives visitors some idea of the current status of a piece of territory and if it is used frequently. Regions under dispute, or used by different animals at different times, may lead to marking battles with every scent marked-over by a new competitor. ==Reproduction behavior==

Estrous cycle and mating Although puppies do not have the urge to procreate, males sometimes engage in sexual play in the form of mounting. In some puppies, this behavior occurs as early as 3 or 4 weeks-of-age. Dogs reach sexual maturity and can reproduce during their first year, in contrast to wolves at two years-of-age. Female dogs 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. Female dogs 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. For several days before estrus, a phase called proestrus, the female dog 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 female dog during proestrus, she may avoid mating by sitting down or turning round and growling or snapping. Estrous behavior in the female dog 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 female dog may also turn, presenting the vulva to the male. 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 female dog for up to 30 to 40 minutes; the penis is twisted 180 degrees in a lateral plane. During this time, prostatic fluid is ejaculated. The female dog can bear another litter within 8 months of the previous one. Dogs are polygamous in contrast to wolves that are generally monogamous. Therefore, dogs have no pair bonding and the protection of a single mate, but rather have multiple mates in a year. 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, one adapted to a life of scavenging instead of hunting. Parenting and early life All of the wild members of the genus Canis display complex coordinated parental behaviors. Wolf pups are cared for primarily by their mother for the first 3 months of their life when she remains in the den with them while they rely on her milk for sustenance and her presence for protection. The father brings her food. Once they leave the den and can chew, the parents and pups from previous years regurgitate food for them. Wolf pups become independent by 5 to 8 months, although they often stay with their parents for years. In contrast, dog pups are cared for by the mother and rely on her for milk and protection but she gets no help from the father nor other dogs. Once pups are weaned around 10 weeks they are independent and receive no further maternal care. ==Behavior problems==

There are many different types of behavioural issues that a dog can exhibit, including growling, snapping, barking, and invading a human's personal space. A survey of 203 dog owners in Melbourne, Australia, found that the main behaviour problems reported by owners were overexcitement (63%) and jumping up on people (56%). Some problems are related to attachment while others are neurological, as seen below. Separation anxiety When dogs are separated from humans, usually the owner, they often display behaviors which can be broken into the following four categories: exploratory behaviour, object play, destructive behaviour, and vocalization, and they are related to the canine's level of arousal. These behaviours may manifest as destructiveness, fecal or urinary elimination, hypersalivation or vocalization among other things. 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. It has been estimated that at least 14% of dogs examined at typical veterinary practices in the United States have shown signs of separation anxiety. Dogs that have been diagnosed with profound separation anxiety can be left alone for no more than a few minutes before they begin to panic and exhibit the behaviors associated with separation anxiety. Separation problems have been found to be linked to the dog's dependency on its owner, not because of disobedience. Environmental management, such as providing chew toys, background sound, or short pre-departure training sessions, can reduce anxiety in mild cases. Calming auditory environments, including specially composed music and soundscapes, have been used effectively to support dogs experiencing separation-related stress. Resource guarding Resource guarding is exhibited by many canines, and is one of the most commonly reported behaviour issues to canine professionals. It is seen when a dog uses specific behaviour patterns so that they can control access to an item, and the patterns are flexible when people are around. If a canine places value on some resource (i.e. food, toys, etc.) they may attempt to guard it from other animals as well as people, which leads to behavioural problems if not treated. The guarding can show in many different ways from rapid ingestion of food to using the body to shield items. It manifests as aggressive behaviour including, but not limited to, growling, barking, or snapping. Some dogs will also resource guard their owners and can become aggressive if the behaviour is allowed to continue. Owners must learn to interpret their dog's body language in order to try to judge the dog's reaction, as visual signals are used (i.e. changes in body posture, facial expression, etc.) to communicate feeling and response. This emotion may feed into other behavioural problems, manifest as attention-seeking behaviour, withdrawing from social activity, or aggression towards their owner or another animal or person. Noise anxiety Canines often fear, and exhibit stress responses to, loud noises. Noise-related anxieties in dogs may be triggered by fireworks, thunderstorms, gunshots, and even loud or sharp bird noises. Associated stimuli may also come to trigger the symptoms of the phobia or anxiety, such as a change in barometric pressure being associated with a thunderstorm, thus causing an anticipatory anxiety. Tail chasing Tail chasing can be classified as a stereotypy. It falls under obsessive compulsive disorder, which is a neuropsychiatric disorder that can present in dogs as canine compulsive disorder. 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. Personality can also play a factor in tail chasing. Dogs who chase their tails have been found to be more shy than those who do not, and some dogs also show a lower level of response during tail chasing bouts. ==Behavior compared to other canids==

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. Compared to wolf and dog pups, golden jackal pups develop aggression at the age of 4–6 weeks when play-fighting frequently escalates into uninhibited biting intended to harm. This aggression ceases by 10–12 weeks when a hierarchy has formed. Tameness Unlike other domestic species which were primarily selected for production-related traits, dogs were initially selected for their behaviors. (i.e. selection for tameness), and emotional processing. 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. Social structure 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. Cognition Despite claims that dogs show more human-like social cognition than wolves, 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. Space Space used by feral dogs is not dissimilar from most other canids in that they use defined traditional areas (home ranges) that tend to be defended against intruders, and have core areas where most of their activities are undertaken. Urban domestic dogs have a home range of 2-61 hectares, in contrast to a feral dog's home range of 58 square kilometers. Wolf home ranges vary from 78 square kilometers where prey is deer, to 2.5 square kilometers at higher latitudes where prey is moose and caribou. Wolves will defend their territory based on prey abundance and pack density, but feral dogs will defend their home ranges all year. Where wolf ranges and feral dog ranges overlap, the feral dogs will site their core areas closer to human settlement. Feral dogs, like their ancestors do participate in pup rearing. Several studies show that feral dogs are not primarily scavengers, despite claims in the popular press. Studies in the modern era show that their diet is very opportunistic, ranging from garbage, carrion to live prey. The primary feature that distinguishes feral from domestic dogs is the degree of reliance or dependence on humans, and in some respect, their behavior toward people. Feral dogs survive and reproduce independently of human intervention or assistance. While it is true that some feral dogs use human garbage for food, others acquire their primary subsistence by hunting and scavenging like other wild canids. Dogs may resort to hunting more than garbage consuming when their garbage food source is scarce. Even well-fed domestic dogs are prone to scavenge; gastro-intestinal veterinary visits increase during warmer weather as dogs are prone to eat decaying material. Some dogs consume feces, which may contain nutrition. On occasion well-fed dogs have been known to scavenge their owners' corpses. ==Dogs in human society==

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. Dogs at work Service dogs are those that are trained to help people with disabilities such as blindness, epilepsy, diabetes and autism. 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. Service dog programs have been established to help individuals suffering from Post Traumatic Stress Disorder (PTSD) and have shown to have positive results. ==Attacks==

The human-dog relationship is based on unconditional trust; however, if this trust is lost it will be difficult to reinstate. 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 was estimated in 2013 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. 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. == Genetic basis of dog behavior ==

It is well established that simple genetic architecture underlies the morphological differences in dog breeds, but the underlying genetic impact of domestic canine behavior is more commonly disputed. Advances in genetic research have found associations between certain behavioral traits and gene regions in domestic dogs, this shines a light on breed specific tendencies or behaviors. Early studies in the genetics of breed specific behaviors in the in herding dogs in the 1940s concluded that 'showing eye' and 'bark' behaviors do not follow simple Mendelian inheritance. Decades later the same conclusion has been reached for all studied behaviors, but the complex modes of inheritance have not been completely deciphered. Like human behavior, canine behavior is a result of the interactions between the protein products coded for by genes, and the environment in which the organism lives. The first study to identify a specific locus associated with a behavioral phenotype in dogs (with genome wide significance) was in 2010, when they found that allelic variation in the cdh2 gene were linked to compulsive behavioral phenotypes. A 2019 genome wide association study concluded that a large proportion of behavioral variance across breeds is attributable to genetic factors. Within breeds, the most heritable traits include characteristics selected for in breeding such as trainability, stranger directed aggression, chasing, and attachment/attention seeking. Another study that compared breed data from C-BARQ, to daily behavioral patterns concluded that separation anxiety and owner directed aggression were the only two out of nine traits not found to have significant heritability. Meanwhile, agitation, attention seeking, barking, excitability, fetching, human/object fear, noise fear, non-owner aggression, and trainability were found to have a genetic basis. Genes containing SNPs associated with dog behavior are likely to be expressed in the brain, contributing to pathways related to the development and expression of behavior and cognition (i.e. they influence behavioral processes through expression in the brain). Examples of chromosome loci for putative SNPs and their associated traits include: For example, attachment and attention seeking behaviors were linked to genes associated with dopamine transport and metabolism. Like some physical diseases, it is conceivable that similar presentation of behavioral traits across breeds could be caused by several different kinds of mutations, and conversely, mutations of the same genes could result in diverse phenotypes. For example, studies have found that certain loci associated breed differences in stranger-directed aggression were associated with SNPs in GRM8, a gene that codes for glutamate receptor, one of the major excitatory neurotransmitters in the central nervous system. Genes associated with temperament and startle response in humans such as OTORD and CACNA1C, were linked to breed differences in fear/fear response. Genes associated with aggression in dogs have been linked to aggressive behavior in humans including CPNE4 and OPCML. Frequency of energetic, boisterous and playful behavior include genes previously linked to resting heart rate, daytime rest, and sleep duration in humans such as TMEM132D, AGMO, SNX29, and CACNA2D3.Trainability has been previously associated with intelligence and information processing speed genes ERG, SNX29, CSMD2, and ATRNL1. == See also ==