Orienting asymmetries and physiological reactivity in dogs' response to human emotional faces.

Recent scientific literature shows that emotional cues conveyed by human vocalizations and odours are processed in an asymmetrical way by the canine brain. In the present study, during feeding behaviour, dogs were suddenly presented with 2-D stimuli depicting human faces expressing the Ekman's six basic emotion (e.g. anger, fear, happiness, sadness, surprise, disgust, and neutral), simultaneously into the left and right visual hemifields. A bias to turn the head towards the left (right hemisphere) rather than the right side was observed with human faces expressing anger, fear, and happiness emotions, but an opposite bias (left hemisphere) was observed with human faces expressing surprise. Furthermore, dogs displayed higher behavioural and cardiac activity to picture of human faces expressing clear arousal emotional state. Overall, results demonstrated that dogs are sensitive to emotional cues conveyed by human faces, supporting the existence of an asymmetrical emotional modulation of the canine brain to process basic human emotions.

Auditory lateralization of conspecific and heterospecific vocalizations in cats.

Auditory lateralization in response to both conspecific and heterospecific vocalizations (dog vocalizations) was observed in 16 tabby cats (Felis catus). Six different vocalizations were used: cat "purring," "meowing" and "growling" and dog typical vocalizations of "disturbance," "isolation" and "play." The head-orienting paradigm showed that cats turned their head with the right ear leading (left hemisphere activation) in response to their typical-species vocalization ("meow" and "purring"); on the other hand, a clear bias in the use of the left ear (right hemisphere activation) was observed in response to vocalizations eliciting intense emotion (dogs' vocalizations of "disturbance" and "isolation"). Overall these findings suggest that auditory sensory domain seems to be lateralized also in cat species, stressing the role of the left hemisphere for intraspecific communication and of the right hemisphere in processing threatening and alarming stimuli.

Right-nostril use during sniffing at arousing stimuli produces higher cardiac activity in jumper horses.

Lateralization in horses, Equus caballus, has been reported at both motor and sensory levels. Here we investigated left- and right-nostril use in 12 jumper horses freely sniffing different emotive stimuli. Results revealed that during sniffing at adrenaline and oestrus mare urine stimuli, horses showed a clear right-nostril bias while just a tendency in the use of the right nostril was observed during sniffing of other odours (food, cotton swab and repellent). Sniffing at adrenaline and urine odours was also accompanied by increasing cardiac activity and behavioural reactivity strengthening the role of the right hemisphere in the analysis of intense emotion and sexual behaviour.

Laterality and performance of agility-trained dogs.

Correlations between lateralised behaviour and performance were investigated in 19 agility-trained dogs (Canis familiaris) by scoring paw preference to hold a food object and relating it to performance during typical agility obstacles (jump/A-frame and weave poles). In addition, because recent behavioural studies reported that visual stimuli of emotional valence presented to one visual hemifield at a time affect visually guided motor responses in dogs, the possibility that the position of the owner respectively in the left and in the right canine visual hemifield might be associated with quality of performance during agility was considered. Dogs' temperament was also measured by an owner-rated questionnaire. The most relevant finding was that agility-trained dogs displayed longer latencies to complete the obstacles with the owner located in their left visual hemifield compared to the right. Interestingly, the results showed that this phenomenon was significantly linked to both dogs' trainability and the strength of paw preference.

Catecholamine plasma levels, IFN-γ serum levels and antibodies production induced by rabies vaccine in dogs selected for their paw preference.

To explore the possible role of the sympathetic nervous activity in the asymmetrical crosstalk between the brain and immune system, catecholamine (E, NE) plasma levels, Interferon-γ (IFN-γ) serum levels and production of antibodies induced by rabies vaccine in dogs selected for their paw preference were measured. The results showed that the direction of behavioural lateralization influenced both epinephrine levels and immune response in dogs. A different kinetic of epinephrine levels after immunization was observed in left-pawed dogs compared to both right-pawed and ambidextrous dogs. The titers of antirabies antibodies were lower in left-pawed dogs than in right-pawed and ambidextrous dogs. Similarly, the IFN-γ serum levels were lower in left-pawed dogs than in the other two groups. Taken together, these findings showed that the left-pawed group appeared to be consistently the different group stressing the fundamental role played by the sympathetic nervous system as a mechanistic basis for the crosstalk between the brain and the immune system.

Detour behaviour in attack-trained dogs: left-turners perform better than right-turners.

Detour behaviour was investigated in attack-trained dogs faced with a "U"-shaped vertical barrier behind which a figurant (target) was located. Left-turners took less time to detour the barrier than right-turners. The most logical explanation for the lateral asymmetries observed in dogs' detour behaviour is to assume that they reflect preferential use of the right or the left eye in visual analysis of the target. Given that the lateral field of each eye of dogs projects mainly to the contralateral side of the brain, shorter latencies to solve the task observed in left-turners (right visual hemifield) with respect to right-turners (left visual hemifield) are consistent with specialisation of the left hemisphere in prey-catching behaviour. Overall our results supported previous evidence that cerebral lateralisation in vertebrates can directly affect visually guided motor responses and have practical implications for personnel involved in the selection of dogs trained specifically to assist police and other law-enforcement personnel in their work.

Effect of Attentional Bias on the 3D Rotated Objects Recognition Ability of Dogs.

The ability to recognize rotated objects has been widely reported in the animal kingdom. Studies on animal and human spatial cognition highlighted the importance of visuo-spatial cognitive capability for surviving in a dynamic world. Although domestic animals are frequently involved in activities requiring a high level of visuo-spatial ability, currently, little is known about their visuo-spatial skills. To investigate this issue, we trained six dogs to discriminate between 3D objects (using a modified version of the Shepard-Metzler task) that were then reproduced digitally on a computer. We found that the dogs recognized three-dimensional objects and their rotated versions (45° and 180°) more easily when presented on the left side of the screen, suggesting right hemisphere superiority in the control of visuo-spatial functions. Moreover, we report inter-individual variability in their performance in the visuo-spatial task. Our preliminary results suggest that dogs could use a rotational invariance process for the discrimination of 3D rotated shapes that deserves further investigation.

Relationship between asymmetric nostril use and human emotional odours in cats.

Cat social behaviour and cognition has received a growing interest during the last decades. Recent studies reported that cats efficiently engage in interspecific communication with humans and suggest that cats are sensitive to human emotional visual and auditory cues. To date, there is no evidence on the social and informative role of human emotional odours, which may affect human-cat communication. In this study, we presented cats with human odours collected in different emotional contexts (fear, happiness, physical stress and neutral) and evaluated the animals' behavioural responses. We found that "fear" odours elicited higher stress levels than "physical stress" and "neutral", suggesting that cats perceived the valence of the information conveyed by "fear" olfactory signals and regulate their behaviour accordingly. Moreover, the prevalent use of the right nostril (right hemisphere activation) with the increase of stress levels, particularly in response to "fear" odours, provides first evidence of lateralized emotional functions of olfactory pathways in cats.

Are temporal features crucial acoustic cues in dog vocal recognition?

To investigate the perceptual mechanisms underlying conspecific vocal recognition in canine species, eighteen dogs were presented with playbacks of normal and reversed versions of typical dog vocalizations. Auditory perception was analysed using the head-turn paradigm, a non-invasive technique extensively employed to study hemispheric specializations for processing conspecific vocalizations in primates. The results revealed that dogs usually turn their heads with the right ear leading (left hemisphere activation) in response to the forward version of their typical calls, and with either no bias and the left ear leading (right hemisphere activation) in response to the reversed call versions. Overall, our findings suggest that temporal features are determinant auditory cues for call sound recognition in dogs, and support earlier findings of the role of the left hemisphere in the analyses of intraspecific communication.

Facial asymmetry in dogs with fear and aggressive behaviors towards humans.

There is now scientific evidence that, in dogs, distinctive facial actions are produced in response to different emotionally-arousing stimuli suggesting a relationship between lateralized facial expressions and emotional states. Although in humans, relationships between facial asymmetry and both emotional and physiological distress have been reported, there are currently no data on the laterality of dogs' facial expressions in response to social stimuli with respect to canine behavioral disorders. The aim of the present work was to investigate the facial asymmetries of dogs with fear and aggressive behavior towards humans during two different emotional situations: (1) while the dogs were alone in the presence of their owners and (2) during the approach of an unfamiliar human being. Overall, our results demonstrated high levels of asymmetries in facial expressions of dogs displaying fear and aggressive behaviors towards humans indicating that measuring facial asymmetries in dogs could prove to be a useful non-invasive tool for investigating physiology-based behavioral disorders.

Motivations of Human Helping Behavior towards Dogs.

Human-dog interactions have a positive effect on human sociality and health. The relationship with dogs helps humans to cope with stress during an emotionally challenging period, such as the COVID-19 pandemic. During this period, a growing global interest in pets has been registered, including the volunteering for shelter/stray dog protection. However, a considerable increase of human dysfunctional interventions toward dogs has been observed in Southern Italy. In this study, we investigated the psychological characteristics of humans volunteering at animal shelter or engaged in stray dog protection. The effect of psychological training and education about dog ethological needs on volunteers' helping behavior was also analyzed. We report that the intervention can improve volunteers' physiological features and, consequently, may enhance human management and dog welfare.

Volumetric assessment of cerebral asymmetries in dogs.

In the present study we quantified volumetric brain asymmetries from computed tomography (CT) scans in 12 healthy dogs, using a semi-automated technique for assessing in vivo structure asymmetry. Volumetric assessment of asymmetrical cerebral lateral ventricle (ALV) was also investigated. Our results showed that seven dogs exhibited a right hemisphere significantly greater than the left, two dogs had a left-greater-than-right hemisphere asymmetry, and finally two dogs displayed no significant brain volumetric asymmetry. This right-biased hemispheric asymmetry supports data reported previously using post-mortem morphological studies in both dogs and other mammalian species.

Emotions and Dog Bites: Could Predatory Attacks Be Triggered by Emotional States?

Dog biting events pose severe public health and animal welfare concerns. They result in several consequences for both humans (including physical and psychological trauma) and the dog involved in the biting episode (abandonment, relocation to shelter and euthanasia). Although numerous epidemiological studies have analyzed the different factors influencing the occurrence of such events, to date the role of emotions in the expression of predatory attacks toward humans has been scarcely investigated. This paper focuses on the influence of emotional states on triggering predatory attacks in dogs, particularly in some breeds whose aggression causes severe consequences to human victims. We suggest that a comprehensive analysis of the dog bite phenomenon should consider the emotional state of biting dogs in order to collect reliable and realistic data about bite episodes.

Odour-Evoked Memory in Dogs: Do Odours Help to Retrieve Memories of Food Location?

The ability of odors to spontaneously trigger specific memories has been widely demonstrated in humans. Although increasing evidence support the role of olfaction on dogs' emotions and cognitive processes, very little research has been conducted on its relationship with memory in this species. The present study aimed at investigating the role of olfaction in the recall of detailed memories originally formed in the presence of a specific odor (i.e., vanilla). To test this, three groups of participants were trained with the same spatial learning task while a specific odor (i.e., vanilla) was dispersed in the testing room. Subjects were then divided in three experimental groups and after 24 h delay, they were presented with the same spatial task. The first group (Group 1) performed the task in the presence of a novel odor (i.e., control), whereas the second (Group 2) and the third group (Group 3) carried out the test in the presence of the vanilla odor and no odor (Group 3), respectively. After a brief delay, the test was presented again to the three groups of dogs: subjects of Group 1 were now tested in the presence of the vanilla odor, whereas the Group 2 was tested with the control odor. The Group 3 received no odor in both tests. A significant improvement of dogs' performance was registered in the control-vanilla odors condition (Group 1), suggesting that the exposure to the odor presented at the encoding time would prompt the recall of spatial memories in dogs.

Emotion Recognition in Cats.

Recent studies demonstrated that cats form social bonds with both conspecifics and humans. One of the key factors regulating social interactions is the transfer of emotions between the individuals. The present study aimed at investigating cats' spontaneous ability to match acoustic and visual signals for the recognition of both conspecific and human emotions. Different conspecific (cat "purr" and "hiss") and heterospecific (human "happiness" and "anger") emotional stimuli were presented to the tested population using a cross-modal paradigm. Results showed that cats are able to cross-modally match pictures of emotional faces with their related vocalizations, particularly for emotions of high intensity. Overall, our findings demonstrate that cats have a general mental representation of the emotions of their social partners, both conspecifics and humans.

Horses associate individual human voices with the valence of past interactions: a behavioural and electrophysiological study.

Brain lateralization is a phenomenon widely reported in the animal kingdom and sensory laterality has been shown to be an indicator of the appraisal of the stimulus valence by an individual. This can prove a useful tool to investigate how animals perceive intra- or hetero-specific signals. The human-animal relationship provides an interesting framework for testing the impact of the valence of interactions on emotional memories. In the present study, we tested whether horses could associate individual human voices with past positive or negative experiences. Both behavioural and electroencephalographic measures allowed examining laterality patterns in addition to the behavioural reactions. The results show that horses reacted to voices associated with past positive experiences with increased attention/arousal (gamma oscillations in the right hemisphere) and indicators of a positive emotional state (left hemisphere activation and ears held forward), and to those associated with past negative experiences with negative affective states (right hemisphere activation and ears held backwards). The responses were further influenced by the animals' management conditions (e.g. box or pasture). Overall, these results, associating brain and behaviour analysis, clearly demonstrate that horses' representation of human voices is modulated by the valence of prior horse-human interactions.

Influence of behavioural lateralization on interleukin-2 and interleukin-6 gene expression in dogs before and after immunization with rabies vaccine.

Evidence in several species, including dogs, has been collected demonstrating that the brain hemispheres modulate the immune system in an asymmetrical way. To study the interactions between immune response and lateralization, three groups of mixed breed dogs were selected on the basis of their performance in a paw preference test involving removal of a piece of sticky tape from the snout. The expression of interleukin-2 (IL-2) and interleukin-6 (IL-6) genes was measured in left-pawed, right-pawed and ambidextrous dogs before and after immunization treatment with a rabies vaccine. The results revealed a relationship between the mRNA expression of IL-2 and IL-6 genes and the direction of behavioural lateralization. Under basal conditions, IL-2 and IL-6 gene expression was higher in left-pawed dogs than in right-pawed and ambidextrous dogs. After the vaccine administration, decreasing levels of IL-2 and IL-6 gene expression were observed in left-pawed and right-pawed dogs, but not in ambidextrous dogs. These findings represent the first evidence that brain lateralization may influence the immune system in dogs by the modulation of mRNA gene expression of cytokines such as IL-2 and IL-6, which have been recognized as key immune-regulatory proteins.

Communication in Dogs.

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.

Lateralized behavior and cardiac activity of dogs in response to human emotional vocalizations.

Over the recent years, the study of emotional functioning has become one of the central issues in dog cognition. Previous studies showed that dogs can recognize different emotions by looking at human faces and can correctly match the human emotional state with a vocalization having a negative emotional valence. However, to this day, little is known about how dogs perceive and process human non-verbal vocalizations having different emotional valence. The current research provides new insights into emotional functioning of the canine brain by studying dogs' lateralized auditory functions (to provide a first insight into the valence dimension) matched with both behavior and physiological measures of arousal (to study the arousal dimension) in response to playbacks related to the Ekman's six basic human emotions. Overall, our results indicate lateralized brain patterns for the processing of human emotional vocalizations, with the prevalent use of the right hemisphere in the analysis of vocalizations with a clear negative emotional valence (i.e. "fear" and "sadness") and the prevalent use of the left hemisphere in the analysis of positive vocalization ("happiness"). Furthermore, both cardiac activity and behavior response support the hypothesis that dogs are sensitive to emotional cues of human vocalizations.

Are dogs red-green colour blind?

Neurobiological and molecular studies suggest a dichromatic colour vision in canine species, which appears to be similar to that of human red-green colour blindness. Here, we show that dogs exhibit a behavioural response similar to that of red-green blind human subjects when tested with a modified version of a test commonly used for the diagnosis of human deuteranopia (i.e. the Ishihara's test). Besides contributing to increasing the knowledge about the perceptual ability of dogs, the present work describes for the first time, to our knowledge, a method that can be used to assess colour vision in the animal kingdom.