Dog-Owner Relationship, Owner Interpretations and Dog Personality Are Connected with the Emotional Reactivity of Dogs.

We evaluated the effect of the dog-owner relationship on dogs' emotional reactivity, quantified with heart rate variability (HRV), behavioral changes, physical activity and dog owner interpretations. Twenty nine adult dogs encountered five different emotional situations (i.e., stroking, a feeding toy, separation from the owner, reunion with the owner, a sudden appearance of a novel object). The results showed that both negative and positive situations provoked signs of heightened arousal in dogs. During negative situations, owners' ratings about the heightened emotional arousal correlated with lower HRV, higher physical activity and more behaviors that typically index arousal and fear. The three factors of The Monash Dog-Owner Relationship Scale (MDORS) were reflected in the dogs' heart rate variability and behaviors: the Emotional Closeness factor was related to increased HRV (p = 0.009), suggesting this aspect is associated with the secure base effect, and the Shared Activities factor showed a trend toward lower HRV (p = 0.067) along with more owner-directed behaviors reflecting attachment related arousal. In contrast, the Perceived Costs factor was related to higher HRV (p = 0.009) along with less fear and less owner-directed behaviors, which may reflect the dog's more independent personality. In conclusion, dogs' emotional reactivity and the dog-owner relationship modulate each other, depending on the aspect of the relationship and dogs' individual responsivity.

Description of movement sensor dataset for dog behavior classification.

Movement sensor data from seven static and dynamic dog behaviors (sitting, standing, lying down, trotting, walking, playing, and (treat) searching i.e. sniffing) was collected from 45 middle to large sized dogs with six degree-of-freedom movement sensors attached to the collar and the harness. With 17 dogs the collection procedure was repeated. The duration of each of the seven behaviors was approximately three minutes. The order of the tasks was varied between the dogs and the two repetitions (for the 17 dogs). The behaviors were annotated post-hoc based on the video recordings made with two camcorders during the tests with one second resolution. The annotations were accurately synchronized with the raw movement sensors data. The annotated data was originally used for training behavior classification machine learning algorithms for classifying the seven behaviors. The developed signal processing and classification algorithms are provided together with the raw measurement data and reference annotations. The description and results of the original investigation that the dataset relates to are found in: P. Kumpulainen, A. Valldeoriola Cardó, S. Somppi, H. Törnqvist, H. Väätäjä, P. Majaranta, Y. Gizatdinova, C. Hoog Antink, V. Surakka, M. V. Kujala, O. Vainio, A. Vehkaoja, Dog behavior classification with movement sensors placed on the harness and the collar, Applied Animal behavior Science, 241 (2021), 105,393.

The Interplay Between Affect, Dog's Physical Activity and Dog-Owner Relationship.

Leaving a dog home alone is part of everyday life for most dog owners. Previous research shows that dog-owner relationship has multifarious effects on dog behavior. However, little is known about the interplay between dog-owner relationship, physical activity of the dog, and affective experiences at the time of the owner leaving home and reunion when the owner comes home. In this paper, we explored how the general (daily, home alone, and over the 2-week study period) physical activity of the dog, and owner's perceptions of the dog's affective state were correlated at those particular moments. Nineteen volunteer dog owners had their dogs (N = 19) wear two activity trackers (ActiGraph wGT2X-GT and FitBark2) for 2 weeks 24 h/day. Prior to the 2-week continuous physical activity measurement period, the owners filled in questionnaires about the dog-owner relationship and the dog behavior. In daily questionnaires, owners described and assessed their own and their perception of the emotion-related experiences of their dog and behavior of the dog at the moment of separation and reunion. The results indicated that the dog-owner relationship has an interplay with the mean daily and weekly physical activity levels of the dog. An indication of strong emotional dog-owner relationship (especially related to the attentiveness of the dog, continuous companionship, and time spent together when relaxing) correlated positively with the mean daily activity levels of the dog during the first measurement week of the study. Results also suggest that the mean daily and over the 2-week measurement period physical activity of the dog correlated the affective experiences of the dog and owner as reported by the owner when the dog was left home alone. More research is needed to understand the interplay between affect, physical activity of the dog, dog-owner relationship, and the effects of these factors on, and their interplay with, the welfare of dogs.

Observing animals and humans: dogs target their gaze to the biological information in natural scenes.

BACKGROUND: This study examines how dogs observe images of natural scenes containing living creatures (wild animals, dogs and humans) recorded with eye gaze tracking. Because dogs have had limited exposure to wild animals in their lives, we also consider the natural novelty of the wild animal images for the dogs. METHODS: The eye gaze of dogs was recorded while they viewed natural images containing dogs, humans, and wild animals. Three categories of images were used: naturalistic landscape images containing single humans or animals, full body images containing a single human or an animal, and full body images containing a pair of humans or animals. The gazing behavior of two dog populations, family and kennel dogs, were compared. RESULTS: As a main effect, dogs gazed at living creatures (object areas) longer than the background areas of the images; heads longer than bodies; heads longer than background areas; and bodies longer than background areas. Dogs gazed less at the object areas vs. the background in landscape images than in the other image categories. Both dog groups also gazed wild animal heads longer than human or dog heads in the images. When viewing single animal and human images, family dogs focused their gaze very prominently on the head areas, but in images containing a pair of animals or humans, they gazed more at the body than the head areas. In kennel dogs, the difference in gazing times of the head and body areas within single or paired images failed to reach significance. DISCUSSION: Dogs focused their gaze on living creatures in all image categories, also detecting them in the natural landscape images. Generally, they also gazed at the biologically informative areas of the images, such as the head, which supports the importance of the head/face area for dogs in obtaining social information. The natural novelty of the species represented in the images as well as the image category affected the gazing behavior of dogs. Furthermore, differences in the gazing strategy between family and kennel dogs was obtained, suggesting an influence of different social living environments and life experiences.

Evaluation of Dry Electrodes in Canine Heart Rate Monitoring.

The functionality of three dry electrocardiogram electrode constructions was evaluated by measuring canine heart rate during four different behaviors: Standing, sitting, lying and walking. The testing was repeated (n = 9) in each of the 36 scenarios with three dogs. Two of the electrodes were constructed with spring-loaded test pins while the third electrode was a molded polymer electrode with Ag/AgCl coating. During the measurement, a specifically designed harness was used to attach the electrodes to the dogs. The performance of the electrodes was evaluated and compared in terms of heartbeat detection coverage. The effect on the respective heart rate coverage was studied by computing the heart rate coverage from the measured electrocardiogram signal using a pattern-matching algorithm to extract the R-peaks and further the beat-to-beat heart rate. The results show that the overall coverage ratios regarding the electrodes varied between 45⁻95% in four different activity modes. The lowest coverage was for lying and walking and the highest was for standing and sitting.

Nasal Oxytocin Treatment Biases Dogs' Visual Attention and Emotional Response toward Positive Human Facial Expressions.

The neuropeptide oxytocin plays a critical role in social behavior and emotion regulation in mammals. The aim of this study was to explore how nasal oxytocin administration affects gazing behavior during emotional perception in domestic dogs. Looking patterns of dogs, as a measure of voluntary attention, were recorded during the viewing of human facial expression photographs. The pupil diameters of dogs were also measured as a physiological index of emotional arousal. In a placebo-controlled within-subjects experimental design, 43 dogs, after having received either oxytocin or placebo (saline) nasal spray treatment, were presented with pictures of unfamiliar male human faces displaying either a happy or an angry expression. We found that, depending on the facial expression, the dogs' gaze patterns were affected selectively by oxytocin treatment. After receiving oxytocin, dogs fixated less often on the eye regions of angry faces and revisited (glanced back at) more often the eye regions of smiling (happy) faces than after the placebo treatment. Furthermore, following the oxytocin treatment dogs fixated and revisited the eyes of happy faces significantly more often than the eyes of angry faces. The analysis of dogs' pupil diameters during viewing of human facial expressions indicated that oxytocin may also have a modulatory effect on dogs' emotional arousal. While subjects' pupil sizes were significantly larger when viewing angry faces than happy faces in the control (placebo treatment) condition, oxytocin treatment not only eliminated this effect but caused an opposite pupil response. Overall, these findings suggest that nasal oxytocin administration selectively changes the allocation of attention and emotional arousal in domestic dogs. Oxytocin has the potential to decrease vigilance toward threatening social stimuli and increase the salience of positive social stimuli thus making eye gaze of friendly human faces more salient for dogs. Our study provides further support for the role of the oxytocinergic system in the social perception abilities of domestic dogs. We propose that oxytocin modulates fundamental emotional processing in dogs through a mechanism that may facilitate communication between humans and dogs.

Human Empathy, Personality and Experience Affect the Emotion Ratings of Dog and Human Facial Expressions.

Facial expressions are important for humans in communicating emotions to the conspecifics and enhancing interpersonal understanding. Many muscles producing facial expressions in humans are also found in domestic dogs, but little is known about how humans perceive dog facial expressions, and which psychological factors influence people's perceptions. Here, we asked 34 observers to rate the valence, arousal, and the six basic emotions (happiness, sadness, surprise, disgust, fear, and anger/aggressiveness) from images of human and dog faces with Pleasant, Neutral and Threatening expressions. We investigated how the subjects' personality (the Big Five Inventory), empathy (Interpersonal Reactivity Index) and experience of dog behavior affect the ratings of dog and human faces. Ratings of both species followed similar general patterns: human subjects classified dog facial expressions from pleasant to threatening very similarly to human facial expressions. Subjects with higher emotional empathy evaluated Threatening faces of both species as more negative in valence and higher in anger/aggressiveness. More empathetic subjects also rated the happiness of Pleasant humans but not dogs higher, and they were quicker in their valence judgments of Pleasant human, Threatening human and Threatening dog faces. Experience with dogs correlated positively with ratings of Pleasant and Neutral dog faces. Personality also had a minor effect on the ratings of Pleasant and Neutral faces in both species. The results imply that humans perceive human and dog facial expression in a similar manner, and the perception of both species is influenced by psychological factors of the evaluators. Especially empathy affects both the speed and intensity of rating dogs' emotional facial expressions.

Dogs Evaluate Threatening Facial Expressions by Their Biological Validity--Evidence from Gazing Patterns.

Appropriate response to companions' emotional signals is important for all social creatures. The emotional expressions of humans and non-human animals have analogies in their form and function, suggesting shared evolutionary roots, but very little is known about how animals other than primates view and process facial expressions. In primates, threat-related facial expressions evoke exceptional viewing patterns compared with neutral or positive stimuli. Here, we explore if domestic dogs (Canis familiaris) have such an attentional bias toward threatening social stimuli and whether observed emotional expressions affect dogs' gaze fixation distribution among the facial features (eyes, midface and mouth). We recorded the voluntary eye gaze of 31 domestic dogs during viewing of facial photographs of humans and dogs with three emotional expressions (threatening, pleasant and neutral). We found that dogs' gaze fixations spread systematically among facial features. The distribution of fixations was altered by the seen expression, but eyes were the most probable targets of the first fixations and gathered longer looking durations than mouth regardless of the viewed expression. The examination of the inner facial features as a whole revealed more pronounced scanning differences among expressions. This suggests that dogs do not base their perception of facial expressions on the viewing of single structures, but the interpretation of the composition formed by eyes, midface and mouth. Dogs evaluated social threat rapidly and this evaluation led to attentional bias, which was dependent on the depicted species: threatening conspecifics' faces evoked heightened attention but threatening human faces instead an avoidance response. We propose that threatening signals carrying differential biological validity are processed via distinctive neurocognitive pathways. Both of these mechanisms may have an adaptive significance for domestic dogs. The findings provide a novel perspective on understanding the processing of emotional expressions and sensitivity to social threat in non-primates.

Comparison of dogs and humans in visual scanning of social interaction.

Previous studies have demonstrated similarities in gazing behaviour of dogs and humans, but comparisons under similar conditions are rare, and little is known about dogs' visual attention to social scenes. Here, we recorded the eye gaze of dogs while they viewed images containing two humans or dogs either interacting socially or facing away: the results were compared with equivalent data measured from humans. Furthermore, we compared the gazing behaviour of two dog and two human populations with different social experiences: family and kennel dogs; dog experts and non-experts. Dogs' gazing behaviour was similar to humans: both species gazed longer at the actors in social interaction than in non-social images. However, humans gazed longer at the actors in dog than human social interaction images, whereas dogs gazed longer at the actors in human than dog social interaction images. Both species also made more saccades between actors in images representing non-conspecifics, which could indicate that processing social interaction of non-conspecifics may be more demanding. Dog experts and non-experts viewed the images very similarly. Kennel dogs viewed images less than family dogs, but otherwise their gazing behaviour did not differ, indicating that the basic processing of social stimuli remains similar regardless of social experiences.

How dogs scan familiar and inverted faces: an eye movement study.

Faces play an important role in communication and identity recognition in social animals. Domestic dogs often respond to human facial cues, but their face processing is weakly understood. In this study, facial inversion effect (deficits in face processing when the image is turned upside down) and responses to personal familiarity were tested using eye movement tracking. A total of 23 pet dogs and eight kennel dogs were compared to establish the effects of life experiences on their scanning behavior. All dogs preferred conspecific faces and showed great interest in the eye area, suggesting that they perceived images representing faces. Dogs fixated at the upright faces as long as the inverted faces, but the eye area of upright faces gathered longer total duration and greater relative fixation duration than the eye area of inverted stimuli, regardless of the species (dog or human) shown in the image. Personally, familiar faces and eyes attracted more fixations than the strange ones, suggesting that dogs are likely to recognize conspecific and human faces in photographs. The results imply that face scanning in dogs is guided not only by the physical properties of images, but also by semantic factors. In conclusion, in a free-viewing task, dogs seem to target their fixations at naturally salient and familiar items. Facial images were generally more attractive for pet dogs than kennel dogs, but living environment did not affect conspecific preference or inversion and familiarity responses, suggesting that the basic mechanisms of face processing in dogs could be hardwired or might develop under limited exposure.

Reactivity of dogs' brain oscillations to visual stimuli measured with non-invasive electroencephalography.

Studying cognition of domestic dogs has gone through a renaissance within the last decades. However, although the behavioral studies of dogs are beginning to be common in the field of animal cognition, the neural events underlying cognition remain unknown. Here, we employed a non-invasive electroencephalography, with adhesive electrodes attached to the top of the skin, to measure brain activity of from 8 domestic dogs (Canis familiaris) while they stayed still to observe photos of dog and human faces. Spontaneous oscillatory activity of the dogs, peaking in the sensors over the parieto-occipital cortex, was suppressed statistically significantly during visual task compared with resting activity at the frequency of 15-30 Hz. Moreover, a stimulus-induced low-frequency (~2-6 Hz) suppression locked to the stimulus onset was evident at the frontal sensors, possibly reflecting a motor rhythm guiding the exploratory eye movements. The results suggest task-related reactivity of the macroscopic oscillatory activity in the dog brain. To our knowledge, the study is the first to reveal non-invasively measured reactivity of brain electrophysiological oscillations in healthy dogs, and it has been based purely on positive operant conditional training, without the need for movement restriction or medication.

Visual event-related potentials of dogs: a non-invasive electroencephalography study.

Previously, social and cognitive abilities of dogs have been studied within behavioral experiments, but the neural processing underlying the cognitive events remains to be clarified. Here, we employed completely non-invasive scalp-electroencephalography in studying the neural correlates of the visual cognition of dogs. We measured visual event-related potentials (ERPs) of eight dogs while they observed images of dog and human faces presented on a computer screen. The dogs were trained to lie still with positive operant conditioning, and they were neither mechanically restrained nor sedated during the measurements. The ERPs corresponding to early visual processing of dogs were detectable at 75-100 ms from the stimulus onset in individual dogs, and the group-level data of the 8 dogs differed significantly from zero bilaterally at around 75 ms at the most posterior sensors. Additionally, we detected differences between the responses to human and dog faces in the posterior sensors at 75-100 ms and in the anterior sensors at 350-400 ms. To our knowledge, this is the first illustration of completely non-invasively measured visual brain responses both in individual dogs and within a group-level study, using ecologically valid visual stimuli. The results of the present study validate the feasibility of non-invasive ERP measurements in studies with dogs, and the study is expected to pave the way for further neurocognitive studies in dogs.

Dogs do look at images: eye tracking in canine cognition research.

Despite intense research on the visual communication of domestic dogs, their cognitive capacities have not yet been explored by eye tracking. The aim of the current study was to expand knowledge on the visual cognition of dogs using contact-free eye movement tracking under conditions where social cueing and associative learning were ruled out. We examined whether dogs spontaneously look at actual objects within pictures and can differentiate between pictures according to their novelty or categorical information content. Eye movements of six domestic dogs were tracked during presentation of digital color images of human faces, dog faces, toys, and alphabetic characters. We found that dogs focused their attention on the informative regions of the images without any task-specific pre-training and their gazing behavior depended on the image category. Dogs preferred the facial images of conspecifics over other categories and fixated on a familiar image longer than on novel stimuli regardless of the category. Dogs' attraction to conspecifics over human faces and inanimate objects might reflect their natural interest, but further studies are needed to establish whether dogs possess picture object recognition. Contact-free eye movement tracking is a promising method for the broader exploration of processes underlying special socio-cognitive skills in dogs previously found in behavioral studies.