In humans, theta (θ) band activity (defined as 4–8 or 5–7 Hz), measured over the frontal midline of the scalp, is an important EEG correlate of voluntary and conscious self-control. Theta waves recorded over the frontal lobes can therefore implicate neocortical control mechanisms. Other likely physiological markers include alpha and beta power, and more recently, a flattened 1/f slope. In the domestic dog, inhibitory self-control is extensively studied, but mostly through behavioural tests. Here, we present a first inquiry into possible EEG correlates of cognitive control in family dogs by comparing short ( 30-second-long) EEG recordings from two conditions: passive wakefulness (baseline) versus a delayed gratification challenge (test). These conditions were recorded in a random sequence within the same session, under overall identical settings. In total, we collected 226 short recordings from fourteen dogs. We observed an increase in neural activity (test > baseline) that resembles human cognitive theta in frequency range (4–8 Hz) and which is similarly more pronounced over the frontal midline. Another important finding is a reduction in broadband power and the flattening of the spectrum, which implicates the work of fronto-parietal connections. While these markers do not differentiate between different facets of cognitive control, our capacity to measure them in awake dogs sets the stage for further EEG-based inquiries into canine cognition.
As the presence of social service robots in everyday settings is increasing, it is important to understand how they shape human behaviour in real-world interactions. We aimed to investigate the behavioural differences shown by visitors towards an autonomously operating non-humanoid service robot and a human server performing the same task—offering candies. Additionally, we compared the behavioural responses to the robot with and without candies and evaluated the efficiency of the robot compared to the human server. Our findings confirmed most of our predictions: visitors were more likely to orient at the robot than the human server, but they were not more likely to withdraw from the approaching robot than from the human server. Visitors showed similar frequencies of orienting, giving way, touching, pointing, and waving toward the robot, regardless of whether it was carrying candies. The robot's tray ran out of candies faster than the human server's in all nine rounds, indicating a high level of robot efficiency and differences in interaction dynamics shown towards the robot and the human server. Together, these findings demonstrate that service robots may elicit distinctive patterns of attention and engagement in public settings, underscoring their capacity to influence human behaviour without inducing avoidance. The results represent a first step towards a more nuanced understanding of human–robot interaction in service contexts and inform the responsible integration of robots into human environments.
Elderly care is a key area of interest in social robotics, yet research on the practical applications and acceptance of such robots in real-world group settings remains limited. In this study, we use an ethorobotic approach to explore the potential of a socially assistive robot (SAR) to elicit interactive behaviours, such as orienting, approaching, withdrawing, touching, talking, and smiling, from elderly residents and other non-resident individuals in a nursing home. The elderly residents demonstrated high levels of acceptance and curiosity toward the robot, whereas most non-resident staff tended to ignore it. Residents showed significantly more interest in the robot than non-residents, and both groups maintained a consistent level of interaction over the 11-day observation period. These exploratory observations suggest that it is indeed possible to design socially interactive robots capable of sustaining long-term engagement with elderly individuals.
Since trainers and handlers typically assess explosives detection dogs (EDDs) without standardised tests, questionnaires can provide valuable insights into expert decision-making. This study aimed to identify preferred breeds and key behavioural traits for explosives detection work by collecting standardised evaluations from EDD trainers and handlers across four European countries. We focused on identifying: i) which breeds are considered best suited for explosives detection work, ii) which behavioural traits are considered most important by experts, iii) how twelve breeds are rated on ten key traits, iv) how these evaluations relate to the breeds currently used in each country and v) how experts’ ratings of the breeds’ olfactory abilities relates to previous experimental finding.Approximately 75% of respondents considered the Malinois and the German shepherd dog (GSD) the best and preferred breeds for explosives detection. Breed-use data differed significantly from expert preferences, with the Malinois being overrepresented (largely due to its high use in France), and the GSD underrepresented (being equally or more common in Portugal and Hungary). Experts’ ratings of the breeds’ olfactory abilities diverged from previous experimental findings; for example, the border collie was rated the lowest by experts, yet it performed well in an olfactory task.In addition to olfactory ability, persistence, willingness to work, and motivation for reward were rated as the most important traits for a successful EDD. The GSD and Belgian shepherd dog (BSD) received the highest scores in both olfactory and non-olfactory traits, whereas the beagle and the basset hound scored lowest on non-olfactory traits.Overall, professionals strive for the optimal combination of availability, cultural habits, olfactory, behavioural, and motivational traits when selecting EDDs, rather than olfactory ability alone. By integrating expert evaluations with previous experimental results, our findings highlight that successful explosives detection performance is perceived to rely on a relatively broad trait profile, which commonly used breeds, such as the Malinois and the GSD, are believed to fulfil the best.
The nasal microbiome may play an important role in dogs’ olfaction, as microbial communities in the nasal cavity can directly interact with the olfactory epithelium. Previous studies have used various sampling methods and depths to examine the canine nasal microbiota and the effect of sampling depth on the detected nasal microbiota remains unclear. To address this, we investigated and compared the nasal microbiota of 81 awake family dogs, sampled at two swabbing depths (15 mm and 25 mm) of the left nostril in the same individuals. The most abundant bacterial taxa in our nasal samples belonged to Moraxella (Gammaproteobacteria), consistent with previous studies. The 15 mm samples had a higher Shannon Diversity Index compared to the 25 mm samples, indicating closer proximity to the external environment. However, we found no significant differences in richness and overall microbiota composition between the two sampling depths. These results indicate that bacterial community profiles within the anterior nasal cavity are similar at the two examined depths. Because the deeper sample was always collected first from the same nostril, potential order effects cannot be excluded. Nevertheless, based on the findings, we recommend using a sampling depth up to 15 mm when examining the nasal microbiota of healthy awake mesocephalic dogs in field settings, as it is less invasive and minimises discomfort.
Investigating prosociality within a comparative framework is crucial for understanding the evolutionary origin and emergence of prosocial behaviours, often considered uniquely human. Thus, in this study, children aged 16-24 months and two distantly related domesticated species, that is, untrained companion dogs and cats, both living in the human niche under relatively similar conditions but characterized by different ecological, evolutionary and developmental backgrounds, were compared. Subjects' spontaneous behaviour in a natural helping situation was assessed, where a familiar human caregiver (parent/owner) searched for an object hidden by the experimenter in the subject's presence without requesting help. In addition, orientation and object-related behaviours were measured, distinguishing those explainable by stimulus enhancement (approaching/manipulating) from those most probably explainable by prosocial behaviours (showing/fetching). To control for motivation, a trial where subjects' favourite treat/toy was hidden was added. All three species showed similar levels of attention towards the caregiver/object. Children and dogs displayed similar levels of object-related behaviour in the test trials, including those explainable by stimulus enhancement and those likely indicative of prosociality. By contrast, cats only displayed showing behaviour (gaze alternation), and even this occurred with a significantly lower probability. In the motivational trial, no species differences were observed in any variable, indicating that cats could also be involved in the problem situation if it was in their own interest. The findings indicate that domestication and close social contact with humans do not, by themselves, induce a tendency towards spontaneous, human-like prosociality. The similar results in dogs and children can be explained by the social/cooperative nature inherited from their ancestors and the specific selection effects that occurred during dog domestication. These factors may have contributed to dogs' exceptional cooperativeness with humans, promoting the development of spontaneous interspecific prosocial tendencies. (c) 2026 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights are reserved, including those for text and data mining, AI training, and similar technologies.
Family dogs’ human-analogue attachment to the owner has long been recognised; however, how this dog–owner attachment develops and changes with age remains an open question. We examined age-related variation and the effect of training experience on the attachment behaviour in family dogs (N = 222) using the Strange Situation Test (SST). Across three age groups (puppy–juvenile, adult, and senior dogs), dogs were assessed by scores for three behavioural factors: attachment to the owner, anxiety in the strange situation, and acceptance of a friendly stranger. Based on three basic behaviours, young puppies under 6 months of age already showed attachment behaviour towards their owners. In puppies–juveniles (2.8–12 months), age was not associated with any of the three factors. In adult dogs (1–7 years), age was positively associated with acceptance scores. In senior dogs (≥ 7 years), attachment scores increased with higher age. Training experience influenced behaviour in adulthood: dogs with greater training experience received lower attachment, higher acceptance, and lower anxiety scores. Further analyses revealed that the association between training and lower attachment was mediated by lower anxiety, likely indicating underactivation of the attachment system in the SST in highly trained dogs. Our findings show that the attachment between dogs and owners develops early, remains fairly consistent through adulthood, and becomes more pronounced in later life as dogs become increasingly dependent on human care. This pattern parallels the increased reliance on attachment figures observed in older humans and highlights family dogs as a unique model for investigating how ageing influences attachment.
The role of sleep in memory consolidation is a widely discussed but still debated area of research. In light of the fact that memory consolidation during sleep is an evolutionary adaptive function, investigating the same phenomenon in nonhuman model species is highly relevant for its understanding. One such species, which has acquired human-analog sociocognitive skills through convergent evolution, is the domestic dog. Family dogs have surfaced as an outstanding animal model in sleep research, and their learning skills (in a social context) are subject to sleep-dependent memory consolidation. These results, however, are correlational, and the next challenge is to establish causality. In the present study, we aimed to adapt a TMR (targeted memory reactivation) paradigm in dogs and investigate its effect on sleep parameters. Dogs (N = 16) learned new commands associated with different locations and afterward took part in a sleep polysomnography recording when they were re-exposed to one of the previously learned commands. The results did not indicate a cueing benefit on choice performance. However, there was evidence for a decrease in choice latency after sleep, while the density (occurrence/minute) of fast sleep spindles was also notably higher during TMR recordings than adaptation recordings from the same animals and even compared with a larger reference sample from a previous work. Our study provides empirical evidence that TMR is feasible with family dogs, even during a daytime nap. Furthermore, the present study highlights several methodological and conceptual challenges for future research.
The family dog is a valid model for studying complex human functions and psychological disorders such as Attention-Deficit/Hyperactivity Disorder (ADHD). Based on prior human research indicating impairments in cognitive flexibility related to ADHD, this study investigates the association between dogs’ ADHD-like traits and reversal learning performance. Since sleep improves learning both in humans and dogs, we also examined its impact in this context. Family dogs (N = 64) completed a two-way choice spatial reversal learning task, followed by a one-hour non-invasive sleep electroencephalography (EEG), and then a second reversal task. We used a validated human analogue questionnaire to assess ADHD. Dogs with higher ADHD scores required more trials to pass the first reversal test, but not after sleep. Electrode application was slower and sleep measurement more likely to fail in dogs with higher ADHD scores. Performance improved more from pre- to post-sleep in high-ADHD dogs if they spent at least 40% of the recording asleep. Our findings align with the human literature showing associations between ADHD and cognitive flexibility in dogs. The main novelty here is the ADHD-related potential benefits of repeated task exposure after sufficient sleep on cognitive flexibility. Cognitive training offers a promising direction to mitigate ADHD-related impairments in dogs.
One of the most striking questions of cognitive ethology is how we got in 50 years from dogs tested as laboratory rats to thick volumes analysing the ethology of family dogs? Due to its shared evolutionary history and developmental environment with humans, the dog was introduced as an inspiring model species of the evolution of human socio-cognitive skills. The human behavioural complex theory provided the conceptual ammunition, and the resulting theoretical framework prioritised the dog, whose social competence was argued to be assembled through convergent evolution, in contrast to homology-based model species. Since then, the dog has been applied as the model of numerous phenomena, such as communicative abilities, attachment, personality, social learning, talent, cooperation/prosocial behaviour, ageing, human psychiatric conditions, language competence, autism spectrum disorder, and attention-deficit/hyperactivity disorder. Recently, behavioural observations could be connected and complemented with genetic and neural research. The dog model became a great success, though an alternative approach argued that dogs were much better represented by free-ranging than companion dogs in research. Still, revealing countless similarities (and differences) between dog and human behaviours/abilities, and especially how dogs integrate these skills, the dog model offered a novel complementary approach to study the evolution of the human mind. It has helped us understand the processes and mental representations underlying dogs’ and humans’ functionally analogous behaviour complexes, and guided how this knowledge can be applied even in future translational research. A better understanding of dogs’ socio-cognitive abilities is also highly beneficial in itself, as our relationship with them has changed. It is no longer sufficient that they have skilfully adapted to our world; we also strive to make our coexistence more comfortable for them.
Most studies on dogs’ olfactory performance focused only on some individuals pre-trained for a task or on specially selected/trained detection dogs. Here, using the Natural Detection Task (NDT) that does not require training, we assessed the effect of several potential factors on the performance of a large sample of dogs (N = 527/tested, 484/analysed). Olfactory success was associated with breeds, but breed groups (selected for olfaction, cooperation, or both) were proven non-relevant, suggesting that breed-specific traits are more influential than functional breed group characteristics. Border collies, selected for herding, reached higher success levels than golden retrievers, Hungarian/German vizslas and basset/bloodhounds, selected for hunting/olfaction tasks. Beagles found the hidden food quicker than border collies, basset/bloodhounds, golden retrievers, Labradors, and cocker spaniels. Dogs with higher Responsiveness to training personality scores were more successful, while higher training level linked to slower successful search. Activity/Excitability scores, attention-deficit/hyperactivity disorder scores, and owners’ rewarding style were not associated with performance. Overall, large within-breed/breed-group variabilities reflect complex interactions between genetic and environmental factors influencing performance. Given the complex nature of life-like detection tasks, the NDT may provide more information about dogs’ olfactory ability than on their potential effectiveness in detection work.
Dogs exhibit natural variability in inattention, hyperactivity and impulsivity traits, sometimes with extreme manifestations resembling to Attention-Deficit/Hyperactivity Disorder (ADHD) symptoms in humans. Using human-standard diagnostic methods, we developed a comprehensive approach for supporting diagnosing ADHD in dogs (N = 1872) based on a validated questionnaire that consists of two parts: a symptom section that provides factor scores for three areas (inattention, hyperactivity and impulsivity), and a functionality section that evaluates the extent to which these symptoms cause functional impairments. Based on the functionality section, we classified dogs as functionally impaired if they showed impairment in at least 4 out of the 7 questions of at least one area (n = 116; 6.2% of the sample). Then, we projected the impaired-non-impaired categories on the ADHD total scores of the symptom section to establish potential cutoff values. The ROC curve analysis resulted in an excellent AUC value (0.861). A cutoff score of 26 was established based on well-defined sensitivity and specificity trade-offs. Finally, we applied the combined thresholds of the two parts of the questionnaire to identify dogs at-risk vs. non-at-risk for ADHD (n = 79; 4.22% of the sample). We offer the first replicable method to screen dogs with suspected ADHD. A final diagnosis could be set after dog-owner pairs participate in relevant behavioural tests and an expert consultation. Allowing for comparisons between ADHD vs. typical groups of family dogs, our method not only facilitates the improvement of the wellbeing of at-risk dogs, but also makes it more feasible to use dogs as a natural model species for studying ADHD.
Self-control is crucial for family dogs in an anthropogenic environment. As valid models of Attention Deficit Hyperactivity Disorder (ADHD), dogs may show behavioural inhibition impairments similar to those seen in humans. This study examines links between dogs' ADHD-like traits and their ability to delay gratification (DG) - a key aspect of self-control impaired in children with ADHD. We tested N = 50 family dogs using an intertemporal choice task, where dogs chose between an immediate low-value reward and a delayed high-value reward. Dogs completed 20 delay trials, with delays increasing by 1 s upon successful waiting. The maximum delay reached and waiting behaviour during successful trials were recorded. ADHD scores were obtained from a validated owner-report questionnaire, and training level was included in the analyses. Higher total ADHD scores were associated with shorter maximum delays, but only in dogs with basic or intermediate training. Inattention scores were negatively linked to DG performance in dogs with basic/intermediate training, while hyperactivity was associated with poorer DG only in dogs with basic training. Impulsivity scores were not linked to success. Although some behavioural consistency was observed during waiting, limited between-individual variation prevented the identification of links with success or ADHD-like traits. Our findings mirror human research, showing links between inattention, hyperactivity, and self-control deficits, supporting dogs' potential as a model to study human ADHD. The lack of association with impulsivity suggests the domain-specificity of this trait. The role of training level highlights its importance as a moderating factor in behavioural studies of self-control.
Gazing toward humans is fundamental in dog-human communication. This study aimed to characterize companion dogs based on their gazing behavior and to explore the potential links between gazing, personality (liveliness, confidence, aggressiveness, and attachment), emotional comfort provided to the owners, and breed. We observed 171 dogs in an ambiguous situation involving a moving stimulus, a remote-controlled toy car and gathered additional data with questionnaires. Our behavior analysis, based on gaze frequency directed at the owner and experimenter, identified four distinct clusters of dogs: low gazers, experimenter-focused gazers, owner-focused gazers, and frequent gazers. We found that experimenter-focused gazers exhibited lower levels of liveliness than frequent gazers and were reported to be less aggressive than low gazers and frequent gazers. Owner-focused gazers provided more emotional comfort to their owners than low gazers. Regarding breeds, 56.5% of German shepherd dogs were low gazers, compared to only 5.3% of golden retrievers. Age, sex, neutered status, and approaching the stimulus were not associated with cluster membership. These results highlight the complexity of dog-human communication, suggesting that personality, emotional factors, and breed shape how dogs seek guidance and support from humans or express anxiety.
Most research into human cognition has focused predominantly on right-handed individuals, ignoring the left-handed minority due to assumed differences in their brain lateralization. Although practical limitations make it challenging to recruit an equal number of left- and right-handed human participants, research into animal models may shed light on the assumed, but rarely tested, associations between motoric handedness, side preference and brain lateralisation. In the present study, we used the family dog, often studied as a model for complex human-analogue socio-cognitive skills. In Study 1, N = 19 dogs completed a range of behavioural tests assessing lateralisation followed by a sleep EEG recording. In Study 2, N = 14 dogs, specifically selected based on prior side bias history in other cognitive experiments, were behaviourally tested in two side-preference tasks and assessed in a sleep EEG measurement. Results show that both negative and positive correlations exist between certain behavioural lateralisation parameters and hemispheric asymmetry in sleep EEG spectrum. These partly support the use-dependent recovery function of sleep (negative correlations) as well as suggest fingerprint-like individual-level associations between sleep and waking behaviour. These findings are relevant for future research capitalizing on the dog as a model for human visuo-spatial attentional biases.
To be able to generalise findings to dogs as a species, many studies use a mixed sample, but applying a single breed sample enables the investigation of several influencing factors without the confounding effect of breed. Here, to reveal the effect of some intrinsic and extrinsic factors on olfactory performance, we investigated the effects of age, personality traits (Activity/Excitability and Responsiveness to training), inattention, hyperactivity and impulsivity, training level, and owner’s rewarding style testing 101 border collies’ performance in the Natural Detection Task. We analysed the dataset of 89 individuals that passed the baseline criteria, while searching for hidden food as a reward at increasing difficulty levels. We found that senior dogs were less successful than late puppies, which is in line with previous findings, where older dogs performed worse than young adults. Some level of training positively affected performance, similar to other studies; even dogs with basic obedience performed better than those with no training, but dogs with advanced or special olfactory training were not superior. Dogs with higher Responsiveness-to-training scores and lower Activity/Excitability scores were more successful. Inattention, hyperactivity, and impulsivity scores and the owner’s general rewarding style did not affect the performance. By focusing on a single breed, instead of a large pool of dogs, we could better identify the influencing effects of different training levels and some personality traits on the olfactory performance than in a mixed sample. Our study highlights the importance of how experience gained in any task situation, as well as certain personality traits, can improve a dog's olfactory performance.
AbstractThe animate-inanimate category distinction is one of the general organizing principles in the primate high-level visual cortex. Much less is known about the visual cortical representations of animacy in non-primate mammals with a different evolutionary trajectory of visual capacities. To compare the functional organization underlying animacy perception of a non-primate to a primate species, here we performed an fMRI study in dogs and humans, investigating how animacy structures neural responses in the visual cortex of the two species. Univariate analyses identified animate-sensitive bilateral occipital and temporal regions, non-overlapping with early visual areas, in both species. Multivariate tests confirmed the categorical representations of animate stimuli in these regions. Regions sensitive to different animate stimulus classes (dog, human, cat) overlapped less in dog than in human brains. Together, these findings reveal that the importance of animate-inanimate distinction is reflected in the organization of higher-level visual cortex, also beyond primates. But a key species difference, that neural representations for animate stimuli are less concentrated in dogs than in humans suggests that certain underlying organizing principles that support the visual perception of animacy in primates may not play a similarly important role in other mammals.
Evidence suggests a link between attention-deficit/hyperactivity disorder (ADHD) and the gut microbiome in humans. Dogs are a valuable model for ADHD research, as their gut microbiome more closely resembles the human gut microbiome in composition and functional overlap compared to rodent models. This study investigated potential associations between dogs’ ADHD-related traits and gut microbiome composition/diversity. We assessed inattention, Hyperactivity, impulsivity, and related functional impairments of 164 family dogs using the validated Dog ADHD and Functional Rating Scale, and analysed bacterial 16S rRNA gene sequences from their faecal samples to assess the gut microbiome composition. Higher relative abundance of members of the family Prevotellaceae and genus Prevotella was associated with lower inattention and inattention-related functional impairment scores by both decision tree and Generalized Linear Model analyses. Hyperactivity, impulsivity, and related functional impairments were found to be moderated by age, suggesting that these traits are predominantly age-related and only secondarily influenced by microbiome composition. ADHD total score was negatively associated with Erysipelotrichaceae and positively associated with Alloprevotella. Dogs with higher functional impairment scores had lower alpha-diversity in their gut microbiome, probably indicating reduced microbial health. This is the first study that found negative associations between inattention and the relative abundance of Prevotellaceae in dogs, which parallels findings in human studies. Our current correlational results in family dogs represent only the first step in gaining more insight into the interplay of gut microbiome and neurodevelopmental processes in non-human animals.
The family dog, in its natural environment, exhibits neuropsychological deficits redolent of human psychiatric disorders, including behaviours that are similar to human attention-deficit/hyperactivity disorder (ADHD) symptoms. Based on standard questionnaire methods in humans, we aimed to develop and validate a detailed, psychometrically improved tool to assess owner views on relevant dog behaviours. We modified available questionnaires by adding items that allow for separate analysis of impulsivity, and items on functional impairment. We collected data from 1168 owners for different validation steps of the new questionnaire and, similarly to assessment of humans where teachers also evaluate as an expert control, we collected data from dog trainers. Exploratory and confirmatory factor analysis revealed 3 factors: inattention (IA), hyperactivity (H) and impulsivity (I), corresponding to all three human symptom dimensions in dogs. Test-retest analyses showed excellent agreement between measurements for all factors. Similarly to findings with humans, trainer-owner rating comparisons showed fair (IA) to moderate (H, I) agreement. As in humans, greater ADHD scores were associated with greater functional impairment scores. We suggest that in dogs, similarly to humans, parallel examination of (extreme) ADHD and functional impairment scores could help distinguish diagnosable individuals, after further validation of the questionnaire using a relevant behaviour test.
Emotionally expressive vocalizations can elicit approach-avoidance responses in humans and non-human animals. We investigated whether artificially generated sounds have similar effects on humans. We assessed whether subjects' reactions were linked to acoustic properties, and associated valence and intensity. We generated 343 artificial sounds with differing call lengths, fundamental frequencies and added acoustic features across 7 categories and 3 levels of biological complexity. We assessed the hypothetical behavioural response using an online questionnaire with a manikin task, in which 172 participants indicated whether they would approach or withdraw from an object emitting the sound. (1) Quieter sounds elicited approach, while loud sounds were associated with avoidance. (2) The effect of pitch was modulated by category, call length and loudness. (2a) Low-pitched sounds in complex sound categories prompted avoidance, while in other categories they elicited approach. (2b) Higher pitch in loud sounds had a distancing effect, while higher pitch in quieter sounds prompted approach. (2c) Longer sounds promoted avoidance, especially at high frequencies. (3) Sounds with higher intensity and negative valence elicited avoidance. We conclude that biologically based acoustic signals can be used to regulate the distance between social robots and humans, which can provide an advantage in interactive scenarios.