Dogs do not use their own experience with novel barriers to infer others' visual access.

Despite extensive research into the Theory of Mind abilities in non-human animals, it remains controversial whether they can attribute mental states to other individuals or whether they merely predict future behaviour based on previous behavioural cues. In the present study, we tested pet dogs (in total, N = 92) on adaptations of the 'goggles test' previously used with human infants and great apes. In both a cooperative and a competitive task, dogs were given direct experience with the properties of novel screens (one opaque, the other transparent) inserted into identical, but differently coloured, tunnels. Dogs learned and remembered the properties of the screens even when, later on, these were no longer directly visible to them. Nevertheless, they were not more likely to follow the experimenter's gaze to a target object when the experimenter could see it through the transparent screen. Further, they did not prefer to steal a forbidden treat first in a location obstructed from the experimenter's view by the opaque screen. Therefore, dogs did not show perspective-taking abilities in this study in which the only available cue to infer others' visual access consisted of the subjects' own previous experience with novel visual barriers. We conclude that the behaviour of our dogs, unlike that of infants and apes in previous studies, does not show evidence of experience-projection abilities.

Functional fixedness in chimpanzees.

Differences in the tool use of non-human primates and humans are subject of ongoing debate. In humans, representations of object functions underpin efficient tool use. Yet, representations of object functions can lead to functional fixedness, which describes the fixation on a familiar tool function leading to less efficient problem solving when the problem requires using the tool for a new function. In the current study, we examined whether chimpanzees exhibit functional fixedness. After solving a problem with a tool, chimpanzees were less efficient in solving another problem which required using the same tool with a different function compared to a control group. This fixation effect was still apparent after a period of nine months and when chimpanzees had learned about the function of a tool by observation of a conspecific. These results suggest that functional fixedness in our closest living relatives likely exists and cast doubt on the notion that stable function representations are uniquely human.

Do dogs preferentially encode the identity of the target object or the location of others' actions?

The ability to make sense of and predict others' actions is foundational for many socio-cognitive abilities. Dogs (Canis familiaris) constitute interesting comparative models for the study of action perception due to their marked sensitivity to human actions. We tested companion dogs (N = 21) in two screen-based eye-tracking experiments, adopting a task previously used with human infants and apes, to assess which aspects of an agent's action dogs consider relevant to the agent's underlying intentions. An agent was shown repeatedly acting upon the same one of two objects, positioned in the same location. We then presented the objects in swapped locations and the agent approached the objects centrally (Experiment 1) or the old object in the new location or the new object in the old location (Experiment 2). Dogs' anticipatory fixations and looking times did not reflect an expectation that agents should have continued approaching the same object nor the same location as witnessed during the brief familiarization phase; this contrasts with some findings with infants and apes, but aligns with findings in younger infants before they have sufficient motor experience with the observed action. However, dogs' pupil dilation and latency to make an anticipatory fixation suggested that, if anything, dogs expected the agents to keep approaching the same location rather than the same object, and their looking times showed sensitivity to the animacy of the agents. We conclude that dogs, lacking motor experience with the observed actions of grasping or kicking performed by a human or inanimate agent, might interpret such actions as directed toward a specific location rather than a specific object. Future research will need to further probe the suitability of anticipatory looking as measure of dogs' socio-cognitive abilities given differences between the visual systems of dogs and primates.

How can I find what I want? Can children, chimpanzees and capuchin monkeys form abstract representations to guide their behavior in a sampling task?

concepts are a powerful tool for making wide-ranging predictions in new situations based on little experience. Whereas looking-time studies suggest an early emergence of this ability in human infancy, other paradigms like the relational match to sample task often fail to detect abstract concepts until late preschool years. Similarly, non-human animals show difficulties and often succeed only after long training regimes. Given the considerable influence of slight task modifications, the conclusiveness of these findings for the development and phylogenetic distribution of abstract reasoning is debated. Here, we tested the abilities of 3 to 5-year-old children, chimpanzees, and capuchin monkeys in a unified and more ecologically valid task design based on the concept of "overhypotheses" (Goodman, 1955). Participants sampled high- and low-valued items from containers that either each offered items of uniform value or a mix of high- and low-valued items. In a test situation, participants should switch away earlier from a container offering low-valued items when they learned that, in general, items within a container are of the same type, but should stay longer if they formed the overhypothesis that containers bear a mix of types. We compared each species' performance to the predictions of a probabilistic hierarchical Bayesian model forming overhypotheses at a first and second level of abstraction, adapted to each species' reward preferences. Children and, to a more limited extent, chimpanzees demonstrated their sensitivity to abstract patterns in the evidence. In contrast, capuchin monkeys did not exhibit conclusive evidence for the ability of abstract knowledge formation.

Investigating belief understanding in children in a nonverbal ambiguous displacement and communication setting.

Finding ways to investigate false belief understanding nonverbally is not just important for preverbal children but also is the only way to assess theory of mind (ToM)-like abilities in nonhuman animals. In this preregistered study, we adapted the design from a previous study on pet dogs to investigate false belief understanding in children and to compare it with belief understanding of those previously tested dogs. A total of 32 preschool children (aged 5-6 years) saw the displacement of a reward and obtained nonverbal cueing of the empty container from an adult communicator holding either a true or false belief. In the false belief condition, when the communicator did not know the location of the reward, children picked the baited container, but not the cued container, more often than the empty one. In the true belief condition, when the communicator witnessed the displacement yet still cued the wrong container, children performed randomly. The children's behavior pattern was at odds with that of the dogs tested in a previous study, which picked the cued container more often when the human communicator held a false belief. In addition to species comparisons, because our task does not require verbal responses or relational sentence understanding, it can also be used in preverbal children. The children in our study behaved in line with the existing ToM literature, whereas most (but not all) dogs from the previously collected sample, although sensitive to differences between the belief conditions, deviated from the children. This difference suggests that using closely matched paradigms and experimental procedures can reveal decisive differences in belief processing between species. It also demonstrates the need for a more comprehensive exploration and direct comparison of the various aspects of false belief processing and ToM in different species to understand the evolution of social cognition.

Dogs Rely On Visual Cues Rather Than On Effector-Specific Movement Representations to Predict Human Action Targets.

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.

Dogs' expectations about occlusion events: from expectancy violation to exploration.

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.

Chimpanzees prepare for alternative possible outcomes.

When facing uncertainty, humans often build mental models of alternative outcomes. Considering diverging scenarios allows agents to respond adaptively to different actual worlds by developing contingency plans (covering one's bases). In a pre-registered experiment, we tested whether chimpanzees (Pan troglodytes) prepare for two mutually exclusive possibilities. Chimpanzees could access two pieces of food, but only if they successfully protected them from a human competitor. In one condition, chimpanzees could be certain about which piece of food the human experimenter would attempt to steal. In a second condition, either one of the food rewards was a potential target of the competitor. We found that chimpanzees were significantly more likely to protect both pieces of food in the second relative to the first condition, raising the possibility that chimpanzees represent and prepare effectively for different possible worlds.

Unwilling or unable? Using three-dimensional tracking to evaluate dogs' reactions to differing human intentions.

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.

Evidence for abstract representations in children but not capuchin monkeys.

The use of abstract higher-level knowledge (also called overhypotheses) allows humans to learn quickly from sparse data and make predictions in new situations. Previous research has suggested that humans may be the only species capable of abstract knowledge formation, but this remains controversial. There is also mixed evidence for when this ability emerges over human development. Kemp et al. (2007) proposed a computational model of how overhypotheses could be learned from sparse examples. We provide the first direct test of this model: an ecologically valid paradigm for testing two species, capuchin monkeys (Sapajus spp.) and 4- to 5-year-old human children. We presented participants with sampled evidence from different containers which suggested that all containers held items of uniform type (type condition) or of uniform size (size condition). Subsequently, we presented two new test containers and an example item from each: a small, high-valued item and a large but low-valued item. Participants could then choose from which test container they would like to receive the next sample - the optimal choice was the container that yielded a large item in the size condition or a high-valued item in the type condition. We compared performance to a priori predictions made by models with and without the capacity to learn overhypotheses. Children's choices were consistent with the model predictions and thus suggest an ability for abstract knowledge formation in the preschool years, whereas monkeys performed at chance level.

Using machine learning to track dogs' exploratory behaviour in the presence and absence of their caregiver.

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.

Eye Tracking in Dogs: Achievements and Challenges.

In this article, we review eye-tracking studies with dogs (Canis familiaris) with a threefold goal; we highlight the achievements in the field of canine perception and cognition using eye tracking, then discuss the challenges that arise in the application of a technology that has been developed in human psychophysics, and finally propose new avenues in dog eye-tracking research. For the first goal, we present studies that investigated dogs' perception of humans, mainly faces, but also hands, gaze, emotions, communicative signals, goal-directed movements, and social interactions, as well as the perception of animations representing possible and impossible physical processes and animacy cues. We then discuss the present challenges of eye tracking with dogs, like doubtful picture-object equivalence, extensive training, small sample sizes, difficult calibration, and artificial stimuli and settings. We suggest possible improvements and solutions for these problems in order to achieve better stimulus and data quality. Finally, we propose the use of dynamic stimuli, pupillometry, arrival time analyses, mobile eye tracking, and combinations with behavioral and neuroimaging methods to further advance canine research and open up new scientific fields in this highly dynamic branch of comparative cognition.

Pupil size changes reveal dogs' sensitivity to motion cues.

Certain motion cues like self-propulsion and speed changes allow human and nonhuman animals to quickly detect animate beings. In the current eye-tracking study, we examined whether dogs' (Canis familiaris) pupil size was influenced by such motion cues. In Experiment 1, dogs watched different videos with normal or reversed playback direction showing a human agent releasing an object. The reversed playback gave the impression that the objects were self-propelled. In Experiment 2, dogs watched videos of a rolling ball that either moved at constant or variable speed. We found that the dogs' pupil size only changed significantly over the course of the videos in the conditions with self-propelled (upward) movements (Experiment 1) or variable speed (Experiment 2). Our findings suggest that dogs orient toward self-propelled stimuli that move at variable speed, which might contribute to their detection of animate beings.

Pet dogs' Behavioural Reaction to Their Caregiver's Interactions with a Third Party: Join in or Interrupt?

Pet dogs are promising candidates to study attachment-related and potentially jealousy-like behaviours in non-human animals, as they form a strong and stable bond with their human caregivers who often engage in affiliative interactions with diverse social partners. Nevertheless, it is still debated whether non-human animals are capable of experiencing such complex emotions. Even though caregivers frequently report observations of jealousy-like behaviours in dogs, behavioural studies in dogs have thus far led to contradictory results. Adding to this complexity, dogs appear extraordinarily skilled in understanding humans' communicative behaviour and can flexibly and diversely interact with them in social contexts. Here, we aimed at investigating (1) whether dogs indeed respond in a jealousy-consistent manner when seeing their caregiver interact in an affiliative way with a remotely controlled, realistic-looking fake dog, or (2) whether they would rather synchronize their reaction to the fake dog with the caregiver's behaviour, or (3) whether they respond directly to the caregiver without paying much attention to the third party. To address what drives the dogs' behaviours in this triadic situation, we compared four groups of dogs who first observed and then joined the interaction of either the caregiver or a stranger greeting or medically examining the fake dog. We found that the dogs initially responded negatively or neutrally when the fake dog entered the room but changed to more positive reactions when the caregiver approached the fake dog, especially if initiating a positive interaction. When being released, more dogs showed friendly behaviours towards the fake dog when the caregiver-rather than the stranger-was interacting with it. At the same time, however, the dogs tried to block the interaction of the caregiver with the fake dog more often than the one of the stranger. In conclusion, we did not find clear evidence for jealousy-like behaviours in dogs during the human-fake dog interactions, but we observed indicators of behavioural synchronization with the caregivers, suggesting that the caregivers' affiliative behaviours directed at a third party may more often facilitate positive than negative interactions in dogs.

The structure of executive functions in preschool children and chimpanzees.

Executive functions (EF) are a core aspect of cognition. Research with adult humans has produced evidence for unity and diversity in the structure of EF. Studies with preschoolers favour a 1-factor model, in which variation in EF tasks is best explained by a single underlying trait on which all EF tasks load. How EF are structured in nonhuman primates remains unknown. This study starts to fill this gap through a comparative, multi-trait multi-method test battery with preschoolers (N = 185) and chimpanzees (N = 55). The battery aimed at measuring working memory updating, inhibition, and attention shifting with three non-verbal tasks per function. For both species the correlations between tasks were low to moderate and not confined to tasks within the same putative function. Factor analyses produced some evidence for the unity of executive functions in both groups, in that our analyses revealed shared variance. However, we could not conclusively distinguish between 1-, 2- or 3-factor models. We discuss the implications of our findings with respect to the ecological validity of current psychometric research.

Inhibitory control and cue relevance modulate chimpanzees' (Pan troglodytes) performance in a spatial foraging task.

Inhibition tasks usually require subjects to exert control to act correctly when a competing action plan is prepotent. In comparative psychology, one concern about the existing inhibition tasks is that the relative contribution of inhibitory control to performance (as compared to learning or object knowledge) is rarely explicitly investigated. We addressed this problem by presenting chimpanzees with a spatial foraging task in which they could acquire food more efficiently by learning which objects were baited. In Experiment 1, we examined how objects that elicited a prepotent approach response, transparent cups containing food, affected their learning rates. Although showing an initial bias to approach these sealed cups with visible food, the chimpanzees learned to avoid them more quickly across sessions compared to a color discrimination. They also learned a color discrimination more quickly if the incorrect cups were sealed such that a piece of food could never be hidden inside them. In Experiment 2, visible food of 2 different types was sealed in the upper part of the cups: 1 type signaled the presence of food reward hidden underneath; the cups with the other type were sealed. The chimpanzees learned more quickly in a congruent condition (the to-be-chosen food cue matched the reward) than in an incongruent condition (the to-be-avoided food cue matched the reward). Together, these findings highlight that performance in inhibition tasks is affected by several other cognitive abilities such as object knowledge, memory, and learning, which need to be quantified before meaningful comparisons can be drawn. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Learning from communication versus observation in great apes.

When human infants are intentionally addressed by others, they tend to interpret the information communicated as being relevant to them and worth acquiring. For humans, this attribution of relevance leads to a preference to learn from communication, making it possible to accumulate knowledge over generations. Great apes are sensitive to communicative cues, but do these cues also activate an expectation of relevance? In an observational learning paradigm, we demonstrated to a sample of nonhuman great apes (bonobos, chimpanzees, orangutans; N = 24) how to operate on a food dispenser device. When apes had the opportunity to choose between an effective and an ineffective method in the baseline conditions, the majority of them chose the effective method. However, when the ineffective method was demonstrated in a communicative way, they failed to prioritize efficiency, even though they were equally attentive in both conditions. This suggests that the ostensive demonstration elicited an expectation of relevance that modified apes' interpretation of the situation, potentially leading to a preference to learn from communication, as human children do.

Dogs' looking times and pupil dilation response reveal expectations about contact causality.

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.

Chimpanzees consider alternative possibilities.

Humans reason not only about actual events (what is), but also about possible events (what could be). Many key operations of human cognition involve the representation of possibilities, including moral judgment, future planning, and causal understanding1. But little is known about the evolutionary roots of this kind of thought. Humans' closest relatives, chimpanzees, possess several cognitive abilities that are closely related to reasoning about alternatives: they plan for the future2, evaluate other's actions3, and reason causally4. However, in the first direct test of the ability to consider alternatives, Redshaw and Suddendorf5 claim that chimpanzees are not able to represent alternative possibilities. Here, using a novel method, we challenge this conclusion: our results suggest that, like human cognition, chimpanzee thought is not limited to what is, but also involves reasoning about what could be the case.