Oxytocin and cortisol are hormones that can influence cognition and behavior, but the relationships between endogenous concentrations and individual differences in cognitive and behavioral phenotypes remain poorly understood. Across mammals, oxytocin has important roles in diverse social behaviors, and in dogs, it has been implicated in human-oriented behaviors such as social gaze and point-following. Cortisol, an end-product of the hypothalamic-pituitary-adrenal (HPA) axis, is often studied in relation to temperament and emotional reactivity, but it is also known to modulate executive functions. In this study, we measured basal fecal cortisol (n = 247) and plasma oxytocin (n = 249) in dog puppies from a pedigreed population (Canine Companions ®). We collected cognitive and behavioral data from these subjects (n = 247), including measures of human-oriented social cognition, memory, inhibitory control, perceptual discriminations, and temperament. Oxytocin concentrations were estimated to be very highly heritable (h2 = 0.90-0.99) and cortisol concentrations were estimated to be moderately-highly heritable (h2 = 0.43-0.47). Bayesian mixed models controlling for relatedness revealed that oxytocin concentrations were positively associated with spatial working memory and displayed a negative quadratic relationship with behavioral laterality, but no credible associations were seen for social measures. Cortisol concentrations exhibited a negative linear relationship with performance on an inhibitory control task and a negative quadratic relationship with bold behavioral reactions to a novel object. Collectively, our results suggest that individual differences in oxytocin and cortisol concentrations are under strong genetic control in dogs and are associated with phenotypic variation in aspects of temperament, behavioral laterality, and executive function.

Machine-learning-based behavioural tracking is a rapidly growing method in the behavioural sciences providing data with high spatial and temporal resolution while reducing the risk of observer bias. Nevertheless, only a few canine behaviour studies have applied this method. In the current study, we used three-dimensional (3D) tracking of the dogs\' bodies to study how separation from the caregiver affected the dogs\' behaviour in a novel environment. During the study, the dogs could move freely in a room equipped with trial-unique objects. We manipulated across trials whether the owner and/or a stranger was present in the room to evaluate the secure base effect, the tendency to explore and play more in the presence of the caregiver compared to another person. This secure base effect is considered a key characteristic of human attachment bonds and has also been described for the dog-caregiver relationship. The tracking data were internally consistent and highly correlated with human scorings and measurements. The results show that both the owner and stranger significantly increased the dogs\' exploration; the dogs also spent more time in the proximity of the owner and stranger location when they were present. Even though the presence of both owner and stranger had a significant effect on the dogs\' behaviour, the effect of the owner was more pronounced. Moreover, in the presence of the stranger the dogs spent more time close to their owner and showed a reduced tail-wagging asymmetry to the right side further supporting the distinct effect of owner and stranger on the dogs\' behaviour. We conclude that machine-driven 3D tracking provides an efficient and reliable access for detailed behavioural analyses of dogs\' exploration and attachment-related behaviours.

Citizen science approaches have grown in popularity over the years, partly due to their ability to reach a wider audience and produce more generalizable samples. In dogs, these studies, though, have been limited in their controls over materials or experimental protocols, with guardians typically reporting results without researcher supervision. Over two studies, we explored and validated a synchronous citizen science approach. We had dog guardians act as experimenters while being supervised by a researcher over Zoom. In study 1, we demonstrated that synchronous citizen science produced equivalent levels of performance to in-lab designs in a choice task. Consistent with past in-lab research, dogs selected a treat (vs. an empty plate) in a two-alternative forced-choice task. In study 2, we showed that Zoom methods are also appropriate for studies utilizing looking time measures. We explored dogs\' looking behaviors when a bag of treats was placed in an unreachable location, and dogs\' guardians were either attentive or inattentive while dogs attempted to retrieve the treats. Consistent with past work, dogs in the attentive condition looked at their guardian for longer periods and had a shorter latency to first look than dogs in the inattentive condition. Overall, we have demonstrated that synchronous citizen science studies with dogs are feasible and produce valid results consistent with those found in a typical lab setting.

Retail dog toys are often provided to companion animals to provide cognitive and physical stimu- lation and improve the animals\' quality of life. These toys, sometimes known as \"enrichment toys\", have been shown to play a role in increasing appetite and activity levels and decreasing undesirable behaviors (e.g., barking, self-isolating behaviors) in some domestic dog (Canis familiaris) populations. In this study, we evaluate the effect of toys on appetite, activity levels, and positive affective states as measures of well- being in companion dogs. Behaviors were compared before and after regular interactions with different types of toys over twelve days. We found that provisioning companion dogs with toys did not significantly alter their activity level, rate of food consumption, or cognitive bias. While dogs who received more complex toys showed a slightly improved cognitive bias, there were no significant differences in behaviors be- tween the subjects who received \"less complex\" toys (e.g., a bone, ball) and \"more complex\" toys (e.g., puzzle toys). We conclude with reflections on the relevance of our subject population to the result seen, and on the different forms of the cognitive bias test.

The ability to predict others\' actions is one of the main pillars of social cognition. We investigated the processes underlying this ability by pitting motor representations of the observed movements against visual familiarity. In two pre-registered eye-tracking experiments, we measured the gaze arrival times of 16 dogs (Canis familiaris) who observed videos of a human or a conspecific executing the same goal-directed actions. On the first trial, when the human agent performed human-typical movements outside dogs\' specific motor repertoire, dogs\' gaze arrived at the target object anticipatorily (i.e., before the human touched the target object). When the agent was a conspecific, dogs\' gaze arrived to the target object reactively (i.e., upon or after touch). When the human agent performed unusual movements more closely related to the dogs\' motor possibilities (e.g., crawling instead of walking), dogs\' gaze arrival times were intermediate between the other two conditions. In a replication experiment, with slightly different stimuli, dogs\' looks to the target object were neither significantly predictive nor reactive, irrespective of the agent. However, when including looks at the target object that were not preceded by looks to the agents, on average dogs looked anticipatorily and sooner at the human agent\'s action target than at the conspecific\'s. Looking times and pupil size analyses suggest that the dogs\' attention was captured more by the dog agent. These results suggest that visual familiarity with the observed action and saliency of the agent had a stronger influence on the dogs\' looking behaviour than effector-specific movement representations in anticipating action targets.

Previous research on human infants has shown that violations of basic physical regularities can stimulate exploration, which may represent a type of hypothesis testing aimed at acquiring knowledge about new causal relationships. In this study, we examined whether a similar connection between expectancy violation and exploration exists in nonhuman animals. Specifically, we investigated how dogs react to expectancy violations in the context of occlusion events. Throughout three experiments, dogs exhibited longer looking times at expectancy-inconsistent events than at consistent ones. This finding was further supported by pupil size analyses in the first two eye-tracking experiments. Our results suggest that dogs expect objects to reappear when they are not obstructed by a screen and consider the size of the occluding screen in relation to the occluded object. In Experiment 3, expectancy violations increased the dogs\' exploration of the target object, similar to the findings with human infants. We conclude that expectancy violations can provide learning opportunities for nonhuman animals as well.

Joint intentionality, the mutual understanding of shared goals or actions to partake in a common task, is considered an essential building block of theory of mind in humans. Domesticated dogs are unusually adept at comprehending human social cues and cooperating with humans, making it possible that they possess behavioral signatures of joint intentionality in interactions with humans. Horschler and colleagues (Anim Behav 183: 159-168, 2022) examined joint intentionality in a service dog population, finding that upon interruption of a joint experience, dogs preferentially re-engaged their former partner over a passive bystander, a behavior argued to be a signature of joint intentionality in human children. In the current study, we aimed to replicate and extend these results in pet dogs. One familiar person played with the dog and then abruptly stopped. We examined if dogs would preferentially re-engage the player instead of a familiar bystander who was also present. Consistent with the findings of Horschler and colleagues (Anim Behav 183: 159-168, 2022), pet dogs preferentially gazed toward and offered the toy to the player significantly more than the familiar bystander. However, no difference was observed in physical contact. These findings provide preliminary evidence for behavioral signatures of joint intentionality in pet dogs, but future work is needed to understand whether this phenomenon extends to other contexts.

The extent to which dogs (Canis familiaris) as a domesticated species understand human intentions is still a matter of debate. The unwilling-unable paradigm has been developed to examine whether nonhuman animals are sensitive to intentions underlying human actions. In this paradigm, subjects tended to wait longer in the testing area when presented with a human that appeared willing but unable to transfer food to them compared to an unwilling (teasing) human. In the present study, we conducted the unwilling-unable paradigm with dogs using a detailed behavioural analysis based on machine-learning driven three-dimensional tracking. Throughout two preregistered experiments, we found evidence, in line with our prediction, that dogs reacted more impatiently to actions signalling unwillingness to transfer food rather than inability. These differences were consistent through two different samples of pet dogs (total n = 96) and they were evident also in the machine-learning generated three-dimensional tracking data. Our results therefore provide robust evidence that dogs distinguish between similar actions (leading to the same outcome) associated with different intentions. However, their reactions did not lead to any measurable preference for one experimenter over the other in a subsequent transfer phase. We discuss different cognitive mechanisms that might underlie dogs\' performance in this paradigm.

UNASSIGNED: The aim of this study was to evaluate the engagement of aging dogs with a cognitively challenging and potentially frustrating task (the impossible task). Based on previous observations, we predicted that dogs showing signs of cognitive impairment in other cognitive tests and owner-completed questionnaires would show reduced engagement with the task.
UNASSIGNED: In this task, dogs were shown a piece of food in a clear container that they could not open; time spent interacting with the container and the experimenter was measured. While the impossible task has not been used as a measure of frustration, the parameters of the test design creates a potential frustrate state, making this assessment appropriate. Thirty-two dogs enrolled in a longitudinal aging study participated in the study. Owners were asked to complete two cognitive dysfunction screening questionnaires (Canine Dementia Scale [CADES] and Canine Cognitive Dysfunction Rating Scale [CCDR]) as well a questionnaire assessing general frustration. Dogs participated in multiple measures of cognitive function as well the impossible task.
UNASSIGNED: Latency to disengage from the impossible task was faster for dogs with higher total (more impaired) CADES (p = 0.02) and total CCDR (p = 0.04) scores. Latency to disengage also correlated with decreased performance in cognitive tests observing social cues (p = 0.01), working memory (p ≤ 0.001), spatial reasoning and reversal learning (p = 0.02), and sustained attention (p = 0.02).
UNASSIGNED: The high correlation with several cognitive measures and the ease of administration of this test makes it a useful tool in evaluating canine cognitive dysfunction syndrome, however it is unclear if increased frustration or other cognitive processes are contributing to the observed changes.

Contact causality is one of the fundamental principles allowing us to make sense of our physical environment. From an early age, humans perceive spatio-temporally contiguous launching events as causal. Surprisingly little is known about causal perception in non-human animals, particularly outside the primate order. Violation-of-expectation paradigms in combination with eye-tracking and pupillometry have been used to study physical expectations in human infants. In the current study, we establish this approach for dogs (Canis familiaris). We presented dogs with realistic three-dimensional animations of launching events with contact (regular launching event) or without contact between the involved objects. In both conditions, the objects moved with the same timing and kinematic properties. The dogs tracked the object movements closely throughout the study but their pupils were larger in the no-contact condition and they looked longer at the object initiating the launch after the no-contact event compared to the contact event. We conclude that dogs have implicit expectations about contact causality.