Pet dogs (Canis familiaris) re-engage humans after joint activity.

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.

Response to Hansen Wheat et al.: Additional analysis further supports the early emergence of cooperative communication in dogs compared to wolves raised with more human exposure.

Here, we address Hansen Wheat et al.'s commentary in this journal in response to Salomons et al. Current Biology, 31(14), 3137-3144.E11, (2021). We conduct additional analyses in response to Hansen Wheat et al.'s two main questions. First, we examine the claim that it was the move to a human home environment which enabled the dog puppies to outperform the wolf puppies in gesture comprehension tasks. We show that the youngest dog puppies who had not yet been individually placed in raisers' homes were still highly skilled, and outperformed similar-aged wolf puppies who had higher levels of human interaction. Second, we address the claim that willingness to approach a stranger can explain the difference between dog and wolf pups' ability to succeed in gesture comprehension tasks. We explain the various controls in the original study that render this explanation insufficient, and demonstrate via model comparison that the covariance of species and temperament also make this parsing impossible. Overall, our additional analyses and considerations support the domestication hypothesis as laid out by Salomons et al. Current Biology, 31(14), 3137-3144.E11, (2021).

Action memory and metamemory.

Actions can enhance memory, exemplified by the enactment effect. In a typical experiment, participants hear a series of simple action phrases (e.g., bounce the ball), which they either carry out (subject-performed tasks, or SPTs), watch the experimenter carry out (experimenter-performed tasks, EPTs), or simply listen to (verbal tasks, VTs). Later memory is usually better for SPTs than for either EPTs or VTs. Although research on action memory is extensive, research on action and metamemory is sparse and produces contradictory results. Furthermore, the metamemory literature has largely ignored the effects of action. Some theoretical perspectives argue that actions produce a particularly effective and automatic form of encoding, and that such nonstrategic encoding should produce inaccurate memory predictions. Other theories argue that action memory relies on executive control processes, suggesting that memory predictions for actions should be just as good (or better) than for control conditions. In Experiments 1a and 1b, participants predicted (with judgements-of-learning, JOLs) whether they would later remember SPTs and EPTs. Resolution (the correlation between JOLs and later recall) was greater for EPTs than SPTs, and not significantly different than zero in the latter case. Experiment 3 produced the same results with SPTs and VTs: resolution was greater for VTs and not significant for SPTs. The results are consistent with nonstrategic accounts of the enactment effect, and also highlight the importance of examining metamemory for actions given that actions can alter metamemory relative to verbal (VT) and other nonaction (EPT) conditions. In addition, the presence of JOLs attenuates the enactment effect, a reactive effect of JOLs similar to that found with other encoding effects. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Cooperative Communication with Humans Evolved to Emerge Early in Domestic Dogs.

Although we know that dogs evolved from wolves, it remains unclear how domestication affected dog cognition. One hypothesis suggests dog domestication altered social maturation by a process of selecting for an attraction to humans.1-3 Under this account, dogs became more flexible in using inherited skills to cooperatively communicate with a new social partner that was previously feared and expressed these unusual social skills early in development.4-6 Here, we comparedog (n = 44) and wolf (n = 37) puppies, 5-18 weeks old, on a battery of temperament and cognition tasks. We find that dog puppies are more attracted to humans, read human gestures more skillfully, and make more eye contact with humans than wolf puppies. The two species are similarly attracted to familiar objects and perform similarly on non-social measures of memory and inhibitory control. These results are consistent with the idea that domestication enhanced the cooperative-communicative abilities of dogs as selection for attraction to humans altered social maturation.

Early-emerging and highly heritable sensitivity to human communication in dogs.

Human cognition is believed to be unique in part because of early-emerging social skills for cooperative communication.1 Comparative studies show that at 2.5 years old, children reason about the physical world similarly to other great apes, yet already possess cognitive skills for cooperative communication far exceeding those in our closest primate relatives.2,3 A growing body of research indicates that domestic dogs exhibit functional similarities to human children in their sensitivity to cooperative-communicative acts. From early in development, dogs flexibly respond to diverse forms of cooperative gestures.4,5 Like human children, dogs are sensitive to ostensive signals marking gestures as communicative, as well as contextual factors needed for inferences about these communicative acts.6-8 However, key questions about potential biological bases for these abilities remain untested. To investigate their developmental and genetic origins, we tested 375 8-week-old dog puppies on a battery of social-cognitive measures. We hypothesized that if dogs' skills for cooperating with humans are biologically prepared, then they should emerge robustly in early development, not require extensive socialization or learning, and exhibit heritable variation. Puppies were highly skillful at using diverse human gestures, and we found no evidence that their performance required learning. Critically, over 40% of the variation in dogs' point-following abilities and attention to human faces was attributable to genetic factors. Our results suggest that these social skills in dogs emerge early in development and are under strong genetic control.

How do non-human primates represent others' awareness of where objects are hidden?

Although non-human primates (NHPs) generally appear to predict how knowledgeable agents use knowledge to guide their behavior, the cognitive mechanisms that enable this remain poorly understood. We assessed the conditions under which NHPs' representations of an agent's awareness break down. Free-ranging rhesus macaques (Macaca mulatta) watched as an agent observed a target object being hidden in one of two boxes. While the agent could no longer see the boxes, the box containing the object flipped open and the object either changed in size/shape (Experiment 1) or color (Experiment 2). Monkeys looked longer when the agent searched for the object incorrectly rather than correctly following the color change (a non-geometric manipulation), but not the size/shape change (a geometric manipulation). Even though the agent maintained knowledge of the object's location in both cases, monkeys no longer expected the agent to search correctly after it had been geometrically (but not non-geometrically) manipulated. Experiment 3 confirmed that monkeys were sensitive to the color manipulation used in Experiment 2, making it unlikely that a failure to perceive the color manipulation accounted for our findings. Our results show that NHPs do not always expect that knowledgeable agents will act on their knowledge to obtain their goals, consistent with heuristic-based accounts of how NHPs represent others' mental states. These findings also suggest that geometric changes that occur outside the agent's perceptual access may disrupt attribution of awareness more so than non-geometric changes.

Dog cognitive development: a longitudinal study across the first 2 years of life.

While our understanding of adult dog cognition has grown considerably over the past 20 years, relatively little is known about the ontogeny of dog cognition. To assess the development and longitudinal stability of cognitive traits in dogs, we administered a battery of tasks to 160 candidate assistance dogs at 2 timepoints. The tasks were designed to measure diverse aspects of cognition, ranging from executive function (e.g., inhibitory control, reversal learning, memory) to sensory discrimination (e.g., vision, audition, olfaction) to social interaction with humans. Subjects first participated as 8-10-week-old puppies, and then were retested on the same tasks at ~ 21 months of age. With few exceptions, task performance improved with age, with the largest effects observed for measures of executive function and social gaze. Results also indicated that individual differences were both early emerging and enduring; for example, social attention to humans, use of human communicative signals, independent persistence at a problem, odor discrimination, and inhibitory control all exhibited moderate levels of rank-order stability between the two timepoints. Using multiple regression, we found that young adult performance on many cognitive tasks could be predicted from a set of cognitive measures collected in early development. Our findings contribute to knowledge about changes in dog cognition across early development as well as the origins and developmental stability of individual differences.

Cognitive characteristics of 8- to 10-week-old assistance dog puppies.

To characterize the early ontogeny of dog cognition, we tested 168 domestic dog, Canis familiaris, puppies (97 females, 71 males; mean age = 9.2 weeks) in a novel test battery based on previous tasks developed and employed with adolescent and adult dogs. Our sample consisted of Labrador retrievers, golden retrievers and Labrador × golden retriever crosses from 65 different litters at Canine Companions for Independence, an organization that breeds, trains and places assistance dogs for people with disabilities. Puppies participated in a 3-day cognitive battery that consisted of 14 tasks measuring different cognitive abilities and temperament traits such as executive function (e.g. inhibitory control, reversal learning, working memory), use of social cues, sensory discriminations and reactivity to and recovery from novel situations. At 8-10 weeks of age, and despite minimal experience with humans, puppies reliably used a variety of cooperative-communicative gestures from humans. Puppies accurately remembered the location of hidden food for delays of up to 20 s, and succeeded in a variety of visual, olfactory and auditory discrimination problems. They also showed some skill at executive function tasks requiring inhibitory control and reversal learning, although they scored lower on these tasks than is typical in adulthood. Taken together, our results confirm the early emergence of sensitivity to human communication in dogs and contextualize these skills within a broad array of other cognitive abilities measured at the same stage of ontogeny.

Do Non-Human Primates Really Represent Others' Beliefs?

Over two decades of research have produced compelling evidence that non-human primates understand some psychological states in other individuals but are unable to represent others' beliefs. Recently, three studies employing anticipatory looking (AL) paradigms reported that non-human primates do show hints of implicitly understanding the beliefs of others. However, measures of AL have been increasingly scrutinized in the human literature owing to extensive replication problems. We argue that new reports of belief representation in non-human primates using AL should be interpreted cautiously because of methodological and theoretical challenges paralleling trends in the human literature. We explore how future work can address these challenges, and conclude by identifying new evolutionary questions raised by the prospect that non-human primates implicitly represent others' beliefs without an explicit belief representation system that guides fitness-relevant behavior.

Leveraging brain-body scaling relationships for comparative studies.

In Horschler et al. (Anim Cognit 22(2):187-198, 2019), we found that two components of executive function (short-term memory and self-control) were strongly associated with estimated absolute brain weight across dog breeds, and argued that dogs present a powerful model for studying evolutionary links between cognition and neuroanatomy due to their extraordinary degree of intraspecific morphological variation. In a commentary on this work, Montgomery (Anim Cognit, 2019) raises concerns about the practice of estimating brain weights from brain-body scaling relationships. Montgomery explores the practical significance of this approach, ultimately concluding that such estimations should be avoided. In this response, we point out some limitations of the analyses presented by Montgomery and consider his conclusions in light of these issues. We then explore the extent to which body weight serves as a valid proxy for brain weight under varying conditions. Through simulations, we show that the consequences of using body weight as a proxy for brain weight depend on parameters including effect size, the correlation between brain and body weight, and the variance in brain and body weight within a sample. Under conditions approximating those in Horschler et al. (Anim Cognit 22(2):187-198, 2019), we find that body weight is a reliable proxy for brain weight, and that statistical results from models using either brain weight or body weight as predictor variables are highly convergent. Nonetheless, we wholeheartedly agree with Montgomery that empirical data on brain weight, structure, and cellular composition will be critical for creating new opportunities to investigate the relationships between neuroanatomy and cognition in dogs.

Do non-human primates really represent others' ignorance? A test of the awareness relations hypothesis.

Non-human primates can often predict how another agent will behave based on that agent's knowledge about the world. But how do non-human primates represent others' knowledge states? Researchers have recently proposed that non-human primates form "awareness relations" to attribute objectively true information to other minds, as opposed to human-like representations that track others' ignorance or false belief states. We present the first explicit test of the awareness relations hypothesis by examining when rhesus macaques' understanding of other agents' knowledge falters. In Experiment 1, monkeys watched an agent observe a piece of fruit (the target object) being hidden in one of two boxes. While the agent's view was occluded, either the fruit moved out of its box and directly back into it, or the box containing the fruit opened and immediately closed. We found that monkeys looked significantly longer when the agent reached incorrectly rather than correctly after the box's movement, but not after the fruit's movement. This result suggests that monkeys did not expect the agent to know the fruit's location when it briefly and arbitrarily moved while the agent could not see it, but did expect the agent to know the fruit's location when only the box moved while the agent could not see it. In Experiment 2, we replicated and extended both findings with a larger sample, a different target object, and opposite directions of motion in the test trials. These findings suggest that monkeys reason about others' knowledge of objects by forming awareness relations which are disrupted by arbitrary spatial manipulation of the target object while an agent has no perceptual access to it.

Absolute brain size predicts dog breed differences in executive function.

Large-scale phylogenetic studies of animal cognition have revealed robust links between absolute brain volume and species differences in executive function. However, past comparative samples have been composed largely of primates, which are characterized by evolutionarily derived neural scaling rules. Therefore, it is currently unknown whether positive associations between brain volume and executive function reflect a broad-scale evolutionary phenomenon, or alternatively, a unique consequence of primate brain evolution. Domestic dogs provide a powerful opportunity for investigating this question due to their close genetic relatedness, but vast intraspecific variation. Using citizen science data on more than 7000 purebred dogs from 74 breeds, and controlling for genetic relatedness between breeds, we identify strong relationships between estimated absolute brain weight and breed differences in cognition. Specifically, larger-brained breeds performed significantly better on measures of short-term memory and self-control. However, the relationships between estimated brain weight and other cognitive measures varied widely, supporting domain-specific accounts of cognitive evolution. Our results suggest that evolutionary increases in brain size are positively associated with taxonomic differences in executive function, even in the absence of primate-like neuroanatomy. These findings also suggest that variation between dog breeds may present a powerful model for investigating correlated changes in neuroanatomy and cognition among closely related taxa.