Dogs' social susceptibility is differentially affected by various dog-Human interactions. A study on family dogs, former shelter dogs and therapy dogs.

When pre-treated with social stimuli prior to testing, dogs are more susceptible to human influence in a food preference task. This means, after a positive social interaction they are more willing to choose the smaller amount of food indicated by the human, as opposed to their baseline preference for the bigger amount. In the current study we investigate if and how various forms of social interaction modulate choices in the same social susceptibility task, testing dogs with varying early life history (pet dogs, therapy dogs, former shelter dogs). In line with previous studies, dogs in general were found to be susceptible to human influence as reflected in the reduced number of "bigger" choices in the human influence, compared to baseline, trials. This was true not only for pet dogs with a normal life history, but also for dogs adopted from a shelter. Therapy dogs, however, did not uniformly change their preference for the bigger quantity of food in the human influence trials; they only did so if prior to testing they had been pre-treated with social stimuli by their owner (but not by a stranger). Pet dogs were also more influenced after pre-treatment with social stimuli by their owner compared to ignoring and separation; however after pre-treatment by a stranger their behaviour did not differ from ignoring and separation. Former shelter dogs on the other hand were equally influenced regardless of pre-treatment by owner versus stranger. In summary these results show that dogs' social susceptibility is modulated by both interactions immediately preceding the test as well as by long term social experiences.

Methodological comparison of cancellation versus two-way choice spatial attention tests in humans and dogs.

Introduction: Behavioural problems in family dogs are amongst the leading reasons for relinquishment to shelters which adversely affects animal welfare. Recent research suggests that certain problematic behavioural patterns might be analogous to human psychiatric disorders. Veterinary diagnosis of such conditions, however, is scarce, probably due to the lack of appropriate measurement tools. The current study focuses on dog behaviour resembling the human hemispatial neglect condition, which manifests itself as a deficit in attention to and awareness of one side of the space. Methods: Healthy human subjects (N = 21) and adult family dogs (N = 23) were tested with tools aimed to measure spatial attention. Tests administered to humans included validated paper and pencil neuropsychological tools to assess hemispatial neglect (cancellation tasks), as well as the canine version of that task (visuo-spatial search task). Dogs were tested with the same visuo-spatial search task as well as a two-way choice task. Results: Results show that both in case of dogs and humans the visuo-spatial search task detects individual variation in subjects' side preferences. However, subjects' performance in the different tasks were not related.

Comment on "Human-directed attachment behaviour in wolves suggests standing ancestral variation for human-dog attachment bonds".

In their recent paper, Hansen Wheat et al. (Ecology and Evolution, 2022, 12, e9299) claimed that hand raised 23-week-old wolves showed the same attachment behaviour towards their handler in the Strange Situation Test (SST) (Determinants of infant behavior, 1969, 4, 111) as dogs. At first glance, their results seem to contradict previous findings that domestication caused a unique change in social-affiliative behaviours in dogs (Animal Behaviour, 2005, 70, 1367). We argue that no persuading evidence was presented to claim that "wolves can show attachment behaviours towards humans comparable to those of dogs". When dealing with a behaviour system (Child Development, 1977, 48, 1184), the subjects' behaviour must meet consistent criteria (Behavioural and Brain Science, 1978, 3, 417), and a few behavioural preferences should not be used to claim the presence of an attachment system, especially, if the experiment violates basic assumptions of the original test. We believe the intriguing scientific question is whether the dog-owner relationship is qualitatively different from what could be observed in the wolf-hand raiser relation. Assessing all available data, our answer is still yes; dogs are unique in this respect.

The influence of voice familiarity and linguistic content on dogs' ability to follow human voice direction.

Domestic dogs are well-known for their abilities to utilize human referential cues for problem solving, including following the direction of human voice. This study investigated whether dogs can locate hidden food relying only on the direction of human voice and whether familiarity with the speaker (owner/stranger) and the relevance of auditory signal features (ostensive addressing indicating the intent for communication to the receiver; linguistic content) affect performance. N = 35 dogs and their owners participated in four conditions in a two-way object choice task. Dogs were presented with referential auditory cues representing different combinations of three contextual parameters: the (I) 'familiarity with the human informant' (owner vs. stranger), the (II) communicative function of attention getter (ostensive addressing vs. non-ostensive cueing) and the (III) 'tone and content of the auditory cue' (high-pitched/potentially relevant vs. low-pitched/potentially irrelevant). Dogs also participated in a 'standard' pointing condition where a visual cue was provided. Significant differences were observed between conditions regarding correct choices and response latencies, suggesting that dogs' response to auditory signals are influenced by the combination of content and intonation of the message and the identity of the speaker. Dogs made correct choices the most frequently when context-relevant auditory information was provided by their owners and showed less success when auditory signals were coming from the experimenter. Correct choices in the 'Pointing' condition were similar to the experimenter auditory conditions, but less frequent compared to the owner condition with potentially relevant auditory information. This was paralleled by shorter response latencies in the owner condition compared to the experimenter conditions, although the two measures were not related. Subjects' performance in response to the owner- and experimenter-given auditory cues were interrelated, but unrelated to responses to pointing gestures, suggesting that dogs' ability to understand the referential nature of auditory cues and visual gestures partly arise from different socio-cognitive skills.

Dog brains are sensitive to infant- and dog-directed prosody.

When addressing preverbal infants and family dogs, people tend to use specific speech styles. While recent studies suggest acoustic parallels between infant- and dog-directed speech, it is unclear whether dogs, like infants, show enhanced neural sensitivity to prosodic aspects of speech directed to them. Using functional magnetic resonance imaging on awake unrestrained dogs we identify two non-primary auditory regions, one that involve the ventralmost part of the left caudal Sylvian gyrus and the temporal pole and the other at the transition of the left caudal and rostral Sylvian gyrus, which respond more to naturalistic dog- and/or infant-directed speech than to adult-directed speech, especially when speak by female speakers. This activity increase is driven by sensitivity to fundamental frequency mean and variance resulting in positive modulatory effects of these acoustic parameters in both aforementioned non-primary auditory regions. These findings show that the dog auditory cortex, similarly to that of human infants, is sensitive to the acoustic properties of speech directed to non-speaking partners. This increased neuronal responsiveness to exaggerated prosody may be one reason why dogs outperform other animals when processing speech.

Six facial prosodic expressions caregivers similarly display to infants and dogs.

Parents tend to use a specific communication style, including specific facial expressions, when speaking to their preverbal infants which has important implications for children's healthy development. In the present study, we investigated these facial prosodic features of caregivers with a novel method that compares infant-, dog- and adult-directed communication. We identified three novel facial displays in addition to the already described three facial expressions (i.e. the 'prosodic faces') that mothers and fathers are typically displaying when interacting with their 1-18 month-old infants and family dogs, but not when interacting with another adult. The so-called Special Happy expression proved to be the most frequent face type during infant- and dog-directed communication which always includes a Duchenne marker to convey an honest and intense happy emotion of the speaker. These results suggest that the 'prosodic faces' play an important role in both adult-infant and human-dog interactions and fulfil specific functions: to call and maintain the partner's attention, to foster emotionally positive interactions, and to strengthen social bonds. Our study highlights the relevance of future comparative studies on facial prosody and its potential contribution to healthy emotional and cognitive development of infants.

Intranasal Oxytocin Improves Social Behavior in Laboratory Beagle Dogs (Canis familiaris) Using a Custom-Made Social Test Battery.

For a long time, oxytocin has been thought to have a generally positive effect on social cognition and prosocial behavior; however, recent results suggested that oxytocin has beneficial effects only under certain conditions. The aim of the present study was to explore potential associations between social competence and the effect of intranasal oxytocin on the social behavior of laboratory beagle dogs. We expected oxytocin treatment to have a more pronounced positive effect on dogs with lower baseline performance in a social test battery. Thirty-six adult dogs of both sexes received 32 IU intranasal oxytocin and physiological saline (placebo) treatment in a double-blind, cross-over design, with 17-20 days between the two sessions. Forty minutes after the treatment, dogs participated in a social test battery consisting of eight situations. The situations were carried out within one session and took 20-30 min to complete. Principal component analysis on the coded behaviors identified four components (Willingness to interact, Preference for social contact, Non-aversive response to nonsocial threat, and Non-aversive response to social threat). The subjects' behavior during the placebo condition was used to assess their baseline performance. We found that oxytocin treatment had a differential effect on the behavior depending on the baseline performance of the individuals in all components, but only two treatment × baseline performance interactions remained significant in a less sensitive analysis. In accordance with our hypothesis, oxytocin administration increased dogs' contact seeking and affiliative behaviors toward humans but only for those with low baseline performance. Dogs with low baseline performance also showed significantly more positive (friendly) reactions to social threat after oxytocin administration than after placebo, while for dogs with high baseline performance, oxytocin administration led to a more negative (fearful) reaction. These results indicate that similar to those on humans, the effects of oxytocin on dogs' social behavior are not universally positive but are constrained by individual characteristics and the context. Nevertheless, oxytocin administration has the potential to improve the social behavior of laboratory beagle dogs that are socially less proficient when interacting with humans, which could have both applied and animal welfare implications.

Social Stimulation by the Owner Increases Dogs' (Canis familiaris) Social Susceptibility in a Food Choice Task-The Possible Effect of Endogenous Oxytocin Release.

Recent evidence suggests a human-like susceptibility to social influence in dogs. For example, dogs tend to ignore their 'natural' preference for the larger amount of food after having seen a human's explicit preference for a smaller quantity. However, it is still unclear whether this tendency to conform to the partner's behaviour can be influenced by social stimuli and/or the neurohormone oxytocin as primers to prosocial predispositions. In Experiment I, eighty two dogs were tested using Prato-Previde et al.'s food quantity preference task. In Experiment I, we investigated in a 2 × 2 design how (i) a 10-minute-long social stimulation by the owner versus a socially ignoring pre-treatment as well as (ii) on-line ostensive communications versus no communication during task demonstration affect dogs' (N = 82) choices in the abovementioned food choice task. Results indicate that the owners' pre-treatment with social stimuli (eye contact, petting) increased dogs' susceptibility to the experimenter's food preference, but the salient ostensive addressing signals accompanying human demonstration masked this social priming effect. In Experiment II, N = 32 dogs from the subjects of Experiment I were retested after oxytocin (OT) or placebo (PL) pre-treatments. This experiment aimed to study whether intranasal administration of oxytocin as compared to placebo treatment would similarly increase dogs' tendency to re-enact the human demonstrator's counterproductive choice in the same task. Results showed an increased susceptibility to the human preference in the OT group, suggesting that both socially stimulating pre-treatment and the intranasal administration of oxytocin have similar priming effects on dogs' social susceptibility.

The implicit reward value of the owner's face for dogs.

It is increasingly assumed that domestication has equipped dogs with unique socio-cognitive skills, which raises the possibility of intriguing parallels between the social motivational systems of the two species. However, the positive incentive value of human facial stimuli is a largely unexplored area. Here, we investigated whether the owner's face serves as a social reinforcer. In a two-way choice task N = 39 dogs were presented with a short video about their owners' head showing the face (facing owner [FO]) vs. the back of the head (non-facing owner). Despite both locations containing equal food reward, dogs approached the container associated with FO more frequently (p < .001), and this was not affected by side, trial order, and choice latency. However, the considerable inter-individual differences in dogs' task performance suggest that the added social component required special social skills which need to be further explored.

Social categorization based on permanent versus transient visual traits in neurotypical children and children with autism spectrum disorder.

The present study was designed to test the relative weight of different types of category markers in children's representations of social and biological kinds. We reasoned that in order to efficiently navigate through the mesh network of overlapping social categories, the representational system dedicated to processing information about social groups should be prepared to flexibly switch between potential ways of categorizing fellow humans. Thus, we hypothesized that children would assign more relevance to transient but symbolic features, such as shirt colour, when categorizing humans than other animal species. Across two experiments, we investigated whether typically developing children as well as children diagnosed with Autism Spectrum Disorder would categorize drawings of humans and dogs along a transient or a biologically set, permanent marker. The results show that both groups of children overwhelmingly selected the permanent feature to categorize dogs, however, they were more likely to categorize fellow humans based on transient features. We suggest that this tendency lays the ground for humans' ability to efficiently represent the complex structure of societies.

Behavioral and Neurophysiological Correlates of Dogs' Individual Sensitivities to Being Observed by Their Owners While Performing a Repetitive Fetching Task.

Ample evidence suggests that dogs possess enhanced skills in reading human visual attention, but it remains to be explored whether they are sensitive to the audience effect in their interactions with humans. The present study aimed to investigate how dogs' behavior is affected by their owners' visual attention while performing a repetitive task (bringing an object back to an unfamiliar experimenter while the owner waited passively). We assumed that if dogs are susceptible to the audience effect, their task persistence and task performance would vary according to their owners' attentiveness. A group of adult pet dogs (N = 27) were repeatedly presented with an object retrieval task by the experimenter (N = 20 trials) while owners either ignored their dogs (Inattentive Owner condition) or paid attention to their dogs' actions (Attentive Owner condition). Behavioral observations were complemented with the owner's reports of their relationships with their dogs (assessed by means of an owner-pet attachment questionnaire) and dogs' spectral EEG sleep profile (recorded during 3-h-long daytime sleep). Although dogs, independently of their owners' attentional state, were generally willing to comply with the fetching task, they were faster to approach the toy object and gazed significantly longer at their owners when he/she was paying attention. This finding is reminiscent of peer influence observed in humans. Further, characteristics of relationship insecurity (relationship anxiety and avoidance) were associated with dogs' task persistence and performance. Dogs of owners with higher relationship anxiety tended to approach the toy object less frequently, and dogs of owners with higher relationship avoidance and anxiety were more hesitant to approach the toy object. We also found that dogs' individual susceptibilities to the audience effect is related to EEG spectral power of both REM and non-REM sleep as well as in pre-sleep (drowsiness) in a trait-like manner. These results, in line with previous findings, support the notion that dogs have a somewhat human-like susceptibility to the audience effect, a trait which might be linked to more complex mechanisms, such as self-presentation or reputation management, helping the two species to become effective social partners.

REM versus Non-REM sleep disturbance specifically affects inter-specific emotion processing in family dogs (Canis familiaris).

Dogs have outstanding capabilities to read human emotional expressions, both vocal and facial. It has also been shown that positively versus negatively valenced dog-human social interactions substantially affect dogs' subsequent sleep. In the present study, we manipulated dogs' (N = 15, in a within subject design) sleep structure by specifically disrupting REM versus Non-REM sleep, while maintaining equal sleep efficiency (monitored via non-invasive polysomnography). We found that both the number of awakenings as well as relative Non-REM (but not relative REM) duration influenced dogs' viewing patterns in a task where sad and happy human faces were simultaneously projected with sad or happy human voice playbacks. In accordance with the emotion laterality hypothesis, the interaction between sound valence and Non-REM sleep duration was specific to images projected to the left (regardless of image-sound congruency). These results reveal the first evidence of a causal link between sleep structure and inter-specific emotion-processing in the family dog.

Comparison of Behavior and Genetic Structure in Populations of Family and Kenneled Beagles.

In dogs, the social and spatial restriction associated with living in a kennel environment could lead to chronic stress and the development of abnormal behaviors ("kennel-dog syndrome"). However, little is known about how kenneled dogs differ from their conspecifics living as pets in human families. In the current study, using a test battery exposing the dogs to novel stimuli, we compared the behavior of three groups of beagles: (1) kenneled dogs living in a restricted environment with limited human contact (N = 78), (2) family dogs living in human families as pets (N = 37), and (3) adopted dogs born in the kenneled population but raised in human families (N = 13). We found one factor comprising most of the test behaviors, labeled as Responsiveness. Family dogs and adopted dogs scored higher in Responsiveness than kenneled dogs. However, 23% of the kenneled dogs were comparable to family and adopted dogs based on a cluster analysis, indicating a similar (positive) reaction to novel stimuli, while 77% of the kenneled dogs were unresponsive (mostly immobile) in at least part of the test. To assess if the behavioral difference between the family and kenneled dogs could be due to genetic divergence of these two populations and/or to lower genetic diversity of the kenneled dogs, we analyzed their genetic structure using 11 microsatellite markers. We found no significant difference between the populations in their genetic diversity (i.e., heterozygosity, level of inbreeding), nor any evidence that the family and kenneled populations originated from different genetic pools. Thus, the behavior difference between the groups more likely reflects a G × E interaction, that is, the influence of specific genetic variants manifesting under specific environmental conditions (kennel life). Nevertheless, some kenneled individuals were (genetically) more resistant to social and environmental deprivation. Selecting for such animals could strongly improve the welfare of kenneled dog populations. Moreover, exploring the genetic background of their higher resilience could also help to better understand the genetics behind stress- and fear-related behaviors.

Reliability of Family Dogs' Sleep Structure Scoring Based on Manual and Automated Sleep Stage Identification.

Non-invasive polysomnography recording on dogs has been claimed to produce data comparable to those for humans regarding sleep macrostructure, EEG spectra and sleep spindles. While functional parallels have been described relating to both affective (e.g., emotion processing) and cognitive (e.g., memory consolidation) domains, methodologically relevant questions about the reliability of sleep stage scoring still need to be addressed. In Study 1, we analyzed the effects of different coders and different numbers of visible EEG channels on the visual scoring of the same polysomnography recordings. The lowest agreement was found between independent coders with different scoring experience using full (3 h-long) recordings of the whole dataset, and the highest agreement within-coder, using only a fraction of the original dataset (randomly selected 100 epochs (i.e., 100 × 20 s long segments)). The identification of drowsiness was found to be the least reliable, while that of non-REM (rapid eye movement, NREM) was the most reliable. Disagreements resulted in no or only moderate differences in macrostructural and spectral variables. Study 2 targeted the task of automated sleep EEG time series classification. Supervised machine learning (ML) models were used to help the manual annotation process by reliably predicting if the dog was sleeping or awake. Logistic regression models (LogREG), gradient boosted trees (GBT) and convolutional neural networks (CNN) were set up and trained for sleep state prediction from already collected and manually annotated EEG data. The evaluation of the individual models suggests that their combination results in the best performance: ~0.9 AUC test scores.

The dog (Canis familiaris) as a translational model of autism: It is high time we move from promise to reality.

Selecting appropriate animal models for a particular human phenomenon is a difficult but important challenge. The difficulty lies in finding animal behaviors that are not only sufficiently relevant and analog to the complex human symptoms (face validity) but also have similar underlying biological and etiological mechanisms (translational or construct validity), and have "human-like" responses to treatment (predictive validity). Over the past several years, the domestic dog (Canis familiaris) has become increasingly proposed as a model for comparative and translational neuroscience. In parallel to the recent advances in canine behavior research, dogs have also been proposed as a model of many human neuropsychiatric conditions, including autism spectrum disorder (ASD). In this opinion paper we will shortly discuss the challenging nature of autism research then summarize the different neurocognitive frameworks for ASD making the case for a canine model of autism. The translational value of a dog model stems from the recognition that (a) there is a large inter-individual variability in the manifestation of dogs' social cognitive abilities including both high and low phenotypic extremes; (b) the phenotypic similarity between the dog and human symptoms are much higher than between the rodent and human symptoms; (c) the symptoms are functionally analogous to the human condition; and (d) more likely to have similar etiology. This article is categorized under: Psychology > Comparative Psychology Cognitive Biology > Evolutionary Roots of Cognition.

Auditory⁻Visual Matching of Conspecifics and Non-Conspecifics by Dogs and Human Infants.

We tested whether dogs and 14⁻16-month-old infants are able to integrate intersensory information when presented with conspecific and heterospecific faces and vocalisations. The looking behaviour of dogs and infants was recorded with a non-invasive eye-tracking technique while they were concurrently presented with a dog and a female human portrait accompanied with acoustic stimuli of female human speech and a dog's bark. Dogs showed evidence of both con- and heterospecific intermodal matching, while infants' looking preferences indicated effective auditory⁻visual matching only when presented with the audio and visual stimuli of the non-conspecifics. The results of the present study provided further evidence that domestic dogs and human infants have similar socio-cognitive skills and highlighted the importance of comparative examinations on intermodal perception.

How do dogs monitor the human's attentional state after challenged by the presence of forbidden food?

INTRODUCTION: Converging evidence suggests that the presence of (attentive) others has a positive effect on people's propensity to conform to social rules. It is also increasingly accepted that pet dogs are promising test subjects to study non-human analogues of 'audience effect.' This study investigates whether dogs show a tendency to change their behavior according to the visual attention of familiar and unfamiliar human partners in a situation in which human partners disallowed the dog from eating a piece of food. METHODS: Dogs (n = 64) participated in two observational conditions (Attentive Owner and Attentive Experimenter) and a control condition in which both human participants engaged in distracting activity. RESULTS: The results showed that the identity of the attentive or inattentive partner has little relevance to the dogs' gazing behavior (i.e., head orientation toward the different partners and the food) and their decisions about breaking or following the rule. This is in line with previous studies suggesting that the presence of the owner predominantly determines the dogs' responses to such situations. DISCUSSION: Further analysis of dogs responding differently to the obedience challenge showed marked differences in the role of the 'audience effect' might play in modulating 'fully obedient,' 'ignorer,' and 'hesitating' dogs' gazing behavior. These findings point to the context-dependent nature of the audience effect in dogs and highlight the importance of frequently ignored individual differences in dogs' tendency to conform to the situational rules.

Similarity between an unfamiliar human and the owner affects dogs' preference for human partner when responding to an unsolvable problem.

This study investigates whether dogs are able to differentiate between people according to whether or not they show similarities to their owners. We hypothesized that dogs would show a preference for the "similar" partner when interacting with unfamiliar humans. After having familiarized with two experimenters displaying different degrees of similarity to their owners, dogs (N = 36) participated in a situation where the desired toy object was made inaccessible in order to find out whether they initiate interaction with the two partners differently. Two different types of "similarity cues" were used (either alone or combined with each other): (1) persistent behavioral characteristics (i.e., familiar vs. strange motion pattern and language usage) and (2) an unfamiliar arbitrary group marker (i.e., one of the potential helpers was wearing clothing similar to that worn by the owner). Results show that although dogs payed equal attention to the human partners displaying various types of similarity to their owners during familiarization, they exhibited a visual attention preference for the human whose motion pattern and language usage were similar to their owner's in the inaccessible-toy task. However, there was weak evidence of discrimination based on the arbitrary group marker (clothing). Although dogs' different tendencies to interact with the potential helpers do not necessarily imply an underlying ability to create social categories based on the degree of similarity between the owner and unfamiliar people, these results suggest that functionally human infant-analogue forms of social categorization may have emerged in dogs.

Dog-Owner Attachment Is Associated With Oxytocin Receptor Gene Polymorphisms in Both Parties. A Comparative Study on Austrian and Hungarian Border Collies.

Variations in human infants' attachment behavior are associated with single nucleotide polymorphisms (SNPs) in the oxytocin receptor (OXTR) gene, suggesting a genetic component to infant-mother attachment. However, due to the genetic relatedness of infants and their mothers, it is difficult to separate the genetic effects of infants' OXTR genotype from the environmental effects of mothers' genotype possibly affecting their parental behavior. The apparent functional analogy between child-parent and dog-owner relationship, however, offers a way to disentangle the effects of these factors because pet dogs are not genetically related to their caregivers. In the present study we investigated whether single nucleotide polymorphisms of pet dogs' OXTR gene (-213AG,-94TC,-74CG) and their owners' OXTR gene (rs53576, rs1042778, rs2254298) are associated with components of dog-owner attachment. In order to investigate whether social-environmental effects modulate the potential genetic influence on attachment, dogs and their owners from two different countries (Austria and Hungary, N = 135 in total) were tested in a modified version of the Ainsworth Strange Situation Test (SST) and questionnaires were also used to collect information about owner personality and attachment style. We coded variables related to three components of attachment behavior in dogs: their sensitivity to the separation from and interaction with the owner (Attachment), stress caused by the unfamiliar environment (Anxiety), and their responsiveness to the stranger (Acceptance). We found that (1) dogs' behavior was significantly associated with polymorphisms in both dogs' and owners' OXTR gene, (2) SNPs in dogs' and owners' OXTR gene interactively influenced dog-human relationship, (3) dogs' attachment behavior was affected by the country of origin, and (4) it was related to their owners' personality as well as attachment style. Thus, the present study provides evidence, for the first time, that both genetic variation in the OXTR gene and various aspects of pet dogs' environmental background are associated with their attachment to their human caregivers.