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Dog Behavior Conference

Communication takes place between members of the same species, as well as between heterospecific individuals, such as the long co-habitation process and inter-dependent relationship present in domestic dogs and humans. Dogs engage in visual communication by modifying different parts of their body; in tactile communication; and also in auditory and olfactory communication, with vocalizations and body odours, respectively. The aim of this review is to provide an overview of the recent literature about dog communication, describing the different nature of the signals used in conspecific and heterospecific interactions and their communicative meaning.

Lateralized dog brain patterns underlying basic neural mechanisms are also discussed, for both conspecific and heterospecific social communication. Dogs have a vast and flexible repertoire of visual, acoustic, and olfactory signals that allow an expressive and fine tuned conspecific and dog—human communication. Dogs use this behavioural repertoire when communicating with humans, employing the same signals used during conspecific interactions, some of which can acquire and carry a different meaning when directed toward humans.

The aim of this review is to provide an overview of the latest progress made in the study of dog communication, describing the different nature of the signals used in conspecific dog—dog and heterospecific dog—human interactions and their communicative meaning. Finally, behavioural asymmetries that reflect lateralized neural patterns involved in both dog—dog and dog—human social communication are discussed.

Communication occurs between members of the same species, as well as between heterospecific individuals, as occurs between domestic dogs and humans [ 1 ]. Living in close contact with humans for at least 30, years [ 2 ], dogs have developed specific skills enabling them to communicate flexibly with humans [ 3 ].

Dogs show a flexible behavioural repertoire when communicating with humans, employing the same signals used in intraspecific interactions dog—dog , some of which can acquire and carry a different meaning when used toward humans e.

They use their whole body to communicate, conveying information intentionally or otherwise [ 8 ]. Not all the signals, in fact, are under voluntary control. When a dog experiences an emotional state, for example anxiety, it releases a specific body odour into the environment [ 8 , 9 ].

Dogs are engaged in visual communication by modifying different parts of their body, in tactile communication, and in auditory and olfactory communication, with vocalizations and body odours, respectively. The aim of this review is to provide an overview on the recent literature about dog communication, describing the different nature of the signals used in conspecific and heterospecific interactions and their communicative meaning.

Dogs communicate visually with other individual modifying the position of different parts of their body see Figure 1 and Figure 2.

For instance, brachycephalic dog lost the flexibility in displaying different facial expressions and dogs with permanently erected ears or with a very short tails lost part of their behavioural repertoire expressed by these anatomical structures [ 10 ]. Therefore, visual communication could be extremely challenging for some dogs, both for correctly delivering and for interpreting visual information. The female is looking at the little red male, asking him to increase the distance.

The little red male is approaching in a curving but conflicting way; he has hackles and his face expresses tension. May be he is testing the reaction of the female, asking her to stand up; the female face expresses threat she probably does not want to interact with him. The two dogs have a very strong relationship. The female is looking at something else with a body language that gives information; she is much more self-confident. Dogs can communicate confidence, alertness, or threat by increasing their body size, pulling themselves up to their full height, and increasing the tension of the body muscles [ 8 ].

On the other hand, dogs can reduce sizes perceived by other individuals by lowering their body and their tail and flattening back their ears to avoid conflicts or during stressful interactions [ 8 , 11 ] Figure 3.

Free-ranging dogs. A The black male displays courtship behavior. His expression shows a closing distance request. A , B The female is showing her intention to avoid a conflict, but also her firm intention to enhance distance to protect her puppy. Dogs wag their tails loosely from side to side to communicate friendliness or their excitability [ 8 ]. Fast movements of the tail, instead, express different inner states according to its position; dogs communicate confidence if they hold their tail high, while a low wagging is generally associated with anxiousness, nervousness, or internal conflict [ 8 , 10 ].

There is now evidence that the direction of tail wagging movements is also directly involved in intraspecific communication. Specifically, when dogs look at stimuli with a positive emotional valence e. The decisive aspect for visual intraspecific communication is that dogs seem to be able to detect tail movement asymmetries of other conspecifics, and thus indirectly deduce their emotional state [ 15 ].

In close-range social interactions, dogs can also obtain and deliver information about their inner state through their facial expression, modifying gaze, ears, and mouth position Figure 3 and Figure 4.

However, recent research has discovered that dogs produce facial expressions as an active attempt to communicate with others [ 16 ]. In this photo two, relaxed facial expressions are shown. The mouths are not tense, the looks are not direct, and the proximity tells us that the two dogs have a good relationship. The eye region plays an important informative role in face recognition in dogs. Dogs usually stare at other individuals to threaten them, while they avoid making eye contact to appease and to decrease the tension during an interaction [ 8 , 10 ].

Eye tracking studies demonstrate, indeed, that dogs address their attention principally to the eye region when processing conspecific faces [ 17 , 18 ]. In this photo, the tension is very high: the Czech wolf is asking the other dog to back off, showing his desire to communicate; he is threatening the white dog, but his look is not directly at the other dog. The white dog instead is much more direct and intense picture taken from a video footage.

Coloured markings around the eyes e. This hypothesis is supported by the fact that there is now clear scientific evidence that, in dogs, colour information may be predominant with respect to brightness [ 19 , 20 ]. Although the mouth region is less investigated compared with the eyes when dogs process conspecific faces, the mouth acquires a particular importance when evaluating whether a facial expression is potentially threatening.

Dogs, indeed, look more at the mouth region of pictures portraying threatening and neutral conspecific facial expressions [ 18 ]. Along with postural and facial displays, dogs can exhibit other behaviours to signal their inner state; for example, they turn their head away from a stimulus when stressed Figure 3 B , they lift their forehead paw to indicate uncertainty, or they lick their lips to communicate their appeasement intentions [ 8 , 21 ].

Dog—human communication has received growing interest over the past twenty years. Dogs, indeed, already show a high sensitivity to human-given cues in an early stage of their development [ 22 , 23 , 24 ], following spontaneously human body postures, gaze direction, and pointing to find a target location [ 25 , 26 , 27 ]. They also prefer to rely more on human gestures rather than auditory cues in a two-choice task, in which the information received is contradictory, suggesting that gestures are more salient for them [ 28 ].

Most importantly, recent studies reported that dogs are skilful in interpreting the communicative intent of humans by understanding the ostensive-referential nature of specific signals, such as eye contact or directed-speech [ 29 , 30 , 31 ].

Furthermore, the flexible comprehension of human gestures allows dogs to efficiently discriminate which of the numerous and different human social behaviours displayed in the everyday life are directed to them [ 7 ]. Dogs, indeed, evaluate the same behaviour differently according to the presence of an ostensive cue that precedes or accompanies it, ignoring the unintended movements [ 29 ].

Among human ostensive signals, eye contact represents the most important and efficient one [ 7 , 29 ]. From an early age, dogs show a spontaneous tendency to gaze at human faces and to make eye contact [ 32 ] in a wide range of contexts, for example, in unsolvable tasks or to beg for food from humans [ 33 , 34 ]. Thus, dogs use eye contact to communicate with humans differently from conspecific communication, in which it represents a clear threatening signal [ 8 ].

On the contrary, in interspecific communication, and in a friendly context, it facilitates the beginning and the maintenance of human—dog interaction [ 35 ].

Therefore, through the domestication process, dogs have modified the functional meaning of this typical behavioural pattern to adapt it to a cross-species communication, acquiring a human-like communication mode [ 30 ]. Furthermore, human—dog mutual gaze enhances the establishment of an affiliative relationship and a social bond between dogs and humans by the same oxytocin-mediated effect described for mother—infant dyad and for human sexual partners [ 5 , 36 , 37 ].

The informative role of the eyes for human—dog communication is also demonstrated by the greater interest by canids in investigating the eye region compared with the other inner facial features in processing human faces [ 17 ]. They use their body position and sustained gaze as a local enhancement signal [ 40 ] or they alternate their gaze between the target object and humans to indicate to them the object location [ 33 , 41 ].

These signals are displayed to communicate with humans and are modulated both by human availability to communicate with them [ 42 ] and by human responses. Dogs, indeed, produce persistently referential signals until they elicit a satisfactory human response [ 43 ], but they are also able to interrupt them when they are no longer successful [ 44 ]. Recent studies have demonstrated the existence of behavioural synchronization between dogs and humans see for review [ 45 ].

The canine synchronizes its locomotor behaviour with that of its owner in different contexts, both indoors [ 46 ] and outdoors [ 47 ], and when facing an unfamiliar human. Thus, authors concluded that, as previously described in humans, this phenomenon increases social cohesion and affiliation in dog—human dyads, contributing to emotional contagion [ 49 ].

Although dogs react to the informative nature of human ostensive-referential cues, they may interpret human gestures as an order rather than understanding the human communicative intent to share information [ 3 , 30 ]. Furthermore, it has been reported in a recent eye-tracking study that dogs are able to discriminate between social and non-social interactions depicted on a picture, showing a longer gaze toward the individuals in a social context compared with a non-social one [ 50 ].

Specifically, domestic dogs show a functional understanding of human emotional facial expressions, responding differently according to its valence. Interestingly, when the informants are inattentive, dogs actively attempt to involve them to obtain information, alternating their gaze between the object and them.

Taken together, these findings demonstrate the existence of social referencing in dogs [ 53 ]. A recent study reports that dogs display mouth-licking behaviour, which is a stress indicator, more often when presented with negative emotional facial expressions compared with positive ones [ 55 ]. Recent scientific literature shows that emotional cues conveyed by human emotional faces are processed in an asymmetrical way by the canine brain. Domestic dogs have a broad and sophisticated vocal repertoire [ 58 ].

Although their vocalizations are similar to their closest relative, the wolf, dogs vocalize in a wider variety of social contexts compared with wolves and they retain this characteristic even into adulthood [ 59 ].

Thus, as described for the foxes, dogs could have acquired a tendency to vocalize more during interactions with humans, which could have been artificially selected, together with other socio-cognitive abilities of understanding human cues. Dogs developed, therefore, novel forms of the pre-existing vocalizations, which acquired different acoustic and functional characteristics, facilitating their communication with humans [ 59 ].

The development of different and numerous vocal signals in dogs could have been modulated, therefore, by their efficacy of conveying specific information to communicate with humans. Among the different vocal signals, the bark is certainly the most typical vocalization of dogs. Contrary to previous beliefs, which claimed that barks are a byproduct of domestication lacking any functional value, recent studies demonstrated the context-related acoustical features of barks [ 60 , 66 , 67 ], suggesting that they are means of communication in dogs.

Barks are short, explosive, and repetitive signals, with a highly variable acoustic structure dominant frequency range between and Hz , differing between breeds and even between individuals [ 60 , 66 ]. Moreover, barking is an allomimetic behaviour, that is, a group activity in which several individuals bark in unison with other conspecifics, mirroring and stimulating each other [ 8 ]. Dog breeds show a different use of barks in their vocal communication.

Wolf-related breeds, for example, Shar-pei, Chow-Chow or Basenji, have a very rare propensity to bark, whereas other breeds present a specific type of barking, such as hunting dogs [ 59 ]. Recent studies report, indeed, that the barks acoustic features vary predictably according to the context; dogs emit longer and lower frequency barks when a stranger approaches them, while high pitched barks are mainly produced in isolation situations [ 63 , 66 ].

Dogs distinguish between the different acoustic structure of barks and react accordingly to its content and the familiarity of the signaler, staying closer to the gate of their house in response to an unfamiliar dog barking at a stranger and remaining inside the house during the barks of a lonely familiar dog [ 67 ].

These findings demonstrate that barks have a functional role in intra-specific communication. Recent studies have reported that, similar to barks, growls also convey meaningful information to dogs [ 62 , 64 , 69 ].

These low-frequency broadband vocalizations are mainly produced during agonistic interactions as a warning or threatening signal or during play interactions [ 8 , 58 ]. Canines can assess the body size of another individual by listening to its growl, correctly matching the sound heard with the picture portraying it [ 69 ]. It has recently been found that canines can extract information about the emotional state of other dogs from their vocalizations.

In fact, they can correctly identify the emotional valence of conspecific vocalizations, associating playful and aggressive ones with the corresponding emotional faces [ 70 ]. Moreover, conspecific vocalizations in the dog brain, as in other vertebrates, are analysed mainly by the left hemisphere, and its involvement depends on the characteristics of the sound. For example, when dogs were presented with the reversed temporal acoustic features of their calls e.

In addition, it is interesting to note that when intraspecific vocalizations elicit intense emotions, a right hemisphere bias appeared, confirming the hypothesis on the role of the right side of the dog brain in the analysis of arousing communicative signals [ 71 ].

Dogs and humans use vocal signals in cross-species communicative interactions that are able to produce changes in other species behaviours [ 72 ].

On one hand, canines understand the meaning of some human words and perceive the emotional content of human vocalizations.

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Enter your mobile number or email address below and we'll send you a link to download the free Kindle App. Then you can start reading Kindle books on your smartphone, tablet, or computer - no Kindle device required. To get the free app, enter your mobile phone number. The answers will surprise and delight you as Alexandra Horowitz A Cognitive Scientist explains how dogs perceive is their daily worlds each other and that other quirky animal. Horowitz introduces the reader to dogs' perceptual and cognitive abilities and then draws a picture of what it might be like to be a dog. What's it like to be able to smell not just every bit of open food in the house but also to smell sadness in humans. Read more Read less.

Account Options Sign in. Top charts. New arrivals. Narrated by Karen White 10 hr 25 min. Switch to the ebook. The bestselling book that asks what dogs know and how they think. The answers will surprise and delight you as Alexandra Horowitz, a cognitive scientist, explains how dogs perceive their daily worlds, each other, and that other quirky animal, the human.

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