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.

Recent developments in parrot cognition: a quadrennial update.

Psittacines, along with corvids, are commonly referred to as 'feathered apes' due to their advanced cognitive abilities. Until rather recently, the research effort on parrot cognition was lagging behind that on corvids, however current developments show that the number of parrot studies is steadily increasing. In 2018, M. L. Lambert et al. provided a comprehensive review on the status of the most important work done so far in parrot and corvid cognition. Nevertheless, only a little more than 4 years after this publication, more than 50 new parrot studies have been published, some of them chartering completely new territory. On the 25th anniversary of Animal Cognition we think this warrants a detailed review of parrot cognition research over the last 4 years. We aim to capture recent developments and current trends in this rapidly expanding and diversifying field.

Do capuchin monkeys (Sapajus apella) use exploration to form intuitions about physical properties?

Humans' flexible innovation relies on our capacity to accurately predict objects' behaviour. These predictions may originate from a "physics-engine" in the brain which simulates our environment. To explore the evolutionary origins of intuitive physics, we investigate whether capuchin monkeys' object exploration supports learning. Two capuchin groups experienced exploration sessions involving multiple copies of two objects, one object was easily opened (functional), the other was not (non-functional). We used two within-subject conditions (enrichment-then-test, and test-only) with two object sets per group. Monkeys then underwent individual test sessions where the objects contained rewards, and they choose one to attempt to open. The monkeys spontaneously explored, performing actions which yielded functional information. At test, both groups chose functional objects above chance. While high performance of the test-only group precluded us from establishing learning during exploration, this study reveals the promise of harnessing primates' natural exploratory tendencies to understand how they see the world.

Do wild raccoons (Procyon lotor) use tools?

Being able to make and use tools was once considered to be an evolutionary hallmark of our species, but has since been documented in other animals. However, for reasons that remain unclear, not all species naturally use tools. Racoons (Procyon lotor) are generalist carnivores that possess many of the physical, cognitive, and behavioural characteristics linked to tool use in other species (e.g. manual dexterity, tactile exploration, relatively large brains, extractive foraging, and sociality). Although raccoons have not been observed using tools outside of experimental captive conditions, wild data involving objective psychometric tests are needed. The current study administered a tool-related task to a wild population of raccoons from 20 locations within the Croatan National Forest, USA. The task required participants to use a stick to extract food from a pipe. To facilitate interpretations of their performances on the task, data were obtained on natural tool availability at the field site and participants' mode of exploring the novel task. None of the participants solved the task despite natural sticks (suitable for solving the task) being widely available across testing locations. Participants were equally likely to smell versus handle novel sticks, which were provided at testing platforms. Limited tactile exploration, but not tool availability, could be at least one factor that reduces these raccoons' opportunities to interact with and learn about novel tools like sticks.

Goffin's cockatoos discriminate objects based on weight alone.

Paying attention to weight is important when deciding upon an object's efficacy or value in various contexts (e.g. tool use, foraging). Proprioceptive discrimination learning, with objects that differ only in weight, has so far been investigated almost exclusively in primate species. Here, we show that while Goffin's cockatoos learn faster when additional colour cues are used, they can also quickly learn to discriminate between objects on the basis of their weight alone. Ultimately, the birds learned to discriminate between visually identical objects on the basis of weight much faster than primates, although methodological differences between tasks should be considered.

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.

Innovative problem solving in macaws.

Behavioural innovations with tool-like objects in non-habitually tool-using species are thought to require complex physical understanding, but the underlying cognitive processes remain poorly understood. A few parrot species are capable of innovating tool-use and borderline tool-use behaviours. We tested this capacity in two species of macaw (Ara ambiguus, n = 9; Ara glaucogularis, n = 8) to investigate if they could solve a problem-solving task through manufacture of a multi-stone construction. Specifically, after having functional experience with a pre-inserted stick tool to push a reward out of a horizontal tube, the subjects were required to insert five stones consecutively from one side to perform the same function as the stick tool with the resulting multi-component construction. One Ara glaucogularis solved the task and innovated the stone construction after the experience with the stick tool. Two more subjects (one of each species) did so after having further functional experience of a single stone pushing a reward out of a shortened tube. These subjects were able to consistently solve the task, but often made errors, for example counter-productive stone insertions from the opposing end, even in some of the successful trials. Conversely, multiple trials without errors also suggested a strong goal direction. Their performance in the follow-up tasks was inconclusive since they sometimes inserted stones into un-baited or blocked 'dummy tubes', but this could have been an attention-deficit behaviour as subjects had not encountered these 'dummy tubes' before. Overall, the successful subjects' performance was so erratic that it proved difficult to conclude whether they had functional understanding of their multi-stone constructions.

String-pulling in the Goffin's cockatoo (Cacatua goffiniana).

Goffin's cockatoos, a parrot species endemic to the Tanimbar Islands in Indonesia, demonstrate remarkable cognitive skills across various technical tasks. These neophilic extractive foragers explore objects with their beak and feet, and are skilled in several modes of tool use. In this study, we confronted the animals for the first time with a vertical string-pulling setup, including a set of classic and novel controls. Nine of the 12 subjects, two of which were subadults, immediately interacted with the single-string task, with seven individuals successfully obtaining the reward on their very first attempt. Four different double string discrimination tests with varying spatial relations were used to assess the Goffin's cockatoos' apprehension of basic physical task properties. We found significant differences in performance between the respective experimental conditions, as well as the development of side biases. The results suggest that while the birds seem to consider simple cause-effect relationships, there is no evidence for a mental representation of the causal mechanisms underlying the string-pulling tasks, as subjects failed the crossed strings condition out of immediate sight. Finally, we provide suggestions on improving the methodology, and discuss our findings in regard to the Goffin's cockatoo's ecology.

Shells as 'extended architecture': to escape isolation, social hermit crabs choose shells with the right external architecture.

Animals' cognitive abilities can be tested by allowing them to choose between alternatives, with only one alternative offering the correct solution to a novel problem. Hermit crabs are evolutionarily specialized to navigate while carrying a shell, with alternative shells representing different forms of 'extended architecture', which effectively change the extent of physical space an individual occupies in the world. It is unknown whether individuals can choose such architecture to solve novel navigational problems. Here, we designed an experiment in which social hermit crabs (Coenobita compressus) had to choose between two alternative shells to solve a novel problem: escaping solitary confinement. Using X-ray microtomography and 3D-printing, we copied preferred shell types and then made artificial alterations to their inner or outer shell architecture, designing only some shells to have the correct architectural fit for escaping the opening of an isolated crab's enclosure. In our 'escape artist' experimental design, crabs had to choose an otherwise less preferred shell, since only this shell had the right external architecture to allow the crab to free itself from isolation. Across multiple experiments, crabs were willing to forgo preferred shells and choose less preferred shells that enabled them to escape, suggesting these animals can solve novel navigational problems with extended architecture. Yet, it remains unclear if individuals solved this problem through trial-and-error or were aware of the deeper connection between escape and exterior shell architecture. Our experiments offer a foundation for further explorations of physical, social, and spatial cognition within the context of extended architecture.

Kangaroos display gazing and gaze alternations during an unsolvable problem task.

Domestication is generally assumed to have resulted in enhanced communication abilities between non-primate mammals and humans, although the number of species studied is very limited (e.g. cats, Felis catus; dogs, Canis familiaris; wolves, Canis lupus; goats, Capra hircus; horses, Equus caballus). In species without hands for pointing, gazing at humans when dealing with inaccessible food during an unsolvable task, and in particular gaze alternations between a human and the unsolvable task (considered forms of showing), are often interpreted as attempts at referential intentional communication. We report that kangaroos, marsupial mammals that have never been domesticated, actively gazed at an experimenter during an unsolvable problem task (10/11 kangaroos tested), thus challenging the notion that this behaviour results from domestication. Nine of the 10 kangaroos additionally showed gaze alternations between the unsolvable task and experimenter. We propose that the potential occurrence of these behaviours displayed towards humans has been underestimated, owing to a narrow focus on domestic animals, as well as a more general eutherian research bias.

Tool-use training temporarily enhances cognitive performance in long-tailed macaques (Macaca fascicularis).

Tool use relies on numerous cognitive functions, including sustained attention and understanding of causality. In this study, we investigated the effects of tool-use training on cognitive performance in primates. Specifically, we applied the Primate Cognition Test Battery to three long-tailed macaques (Macaca fascicularis) at different stages of a training procedure that consisted of using a rake to retrieve out-of-reach food items. In addition, we evaluated a control group (n = 3) performing a grasping task, in order to account for possible effects related to a simple motor act. Our results showed that tool-use training enhances mean performance in the physical cognition domain, i.e. the understanding of spatial relations, numerosity and causality. In particular, causal cognition (evaluating noise- and shape-related causality and understanding of tool properties) showed significant improvement after training, whereas spatial cognition (evaluating spatial memory, object permanence, rotation and transposition) showed a trend to improvement. Despite these findings, none of our trained monkeys succeeded in the tool-use task of the Primate Cognition Test Battery, which involved an unfamiliar tool. Some training-related effects did not persist after a 35-day resting period, suggesting that continuous practice may be necessary, or that a longer training period before resting may be needed to better maintain cognitive performance. In contrast with the training group, the control group did not display any change in cognitive performance. This finding paves the way to further investigation into the link between tool-use behaviour and the evolution of primate cognition.

Practice makes perfect: familiarity of task determines success in solvable tasks for free-ranging dogs (Canis lupus familiaris).

Domestic dogs' (Canis lupus familiaris) socio-cognitive faculties have made them highly sensitive to human social cues. While dogs often excel at understanding human communicative gestures, they perform comparatively poorly in problem-solving and physical reasoning tasks. This difference in their behaviour could be due to the lifestyle and intense socialization, where problem solving and physical cognition are less important than social cognition. Free-ranging dogs live in human-dominated environments, not under human supervision and are less socialized. Being scavengers, they often encounter challenges where problem solving is required in order to get access to food. We tested Indian street dogs in familiar and unfamiliar independent solvable tasks and quantified their persistence and dependence on a novel human experimenter, in addition to their success in solving a task. Our results indicate that free-ranging dogs succeeded and persisted more in the familiar task as compared to the unfamiliar one. They showed negligible amount of human dependence in the familiar task, but showed prolonged gazing and considerable begging behaviour to the human experimenter in the context of the unfamiliar task. Cognitive abilities of free-ranging dogs thus play a pivotal role in determining task-associated behaviours based on familiarity. In addition to that, these dogs inherently tend to socialize with and depend on humans, even if they are strangers. Our results also illustrate free-ranging dogs' low competence at physical cognitive tasks.

Social learning in nest-building birds: a role for familiarity.

It is becoming apparent that birds learn from their own experiences of nest building. What is not clear is whether birds can learn from watching conspecifics build. As social learning allows an animal to gain information without engaging in costly trial-and-error learning, first-time builders should exploit the successful habits of experienced builders. We presented first-time nest-building male zebra finches with either a familiar or an unfamiliar conspecific male building with material of a colour the observer did not like. When given the opportunity to build, males that had watched a familiar male build switched their material preference to that used by the familiar male. Males that observed unfamiliar birds did not. Thus, first-time nest builders use social information and copy the nest material choices when demonstrators are familiar but not when they are strangers. The relationships between individuals therefore influence how nest-building expertise is socially transmitted in zebra finches.

Young children spontaneously invent wild great apes' tool-use behaviours.

The variety and complexity of human-made tools are unique in the animal kingdom. Research investigating why human tool use is special has focused on the role of social learning: while non-human great apes acquire tool-use behaviours mostly by individual (re-)inventions, modern humans use imitation and teaching to accumulate innovations over time. However, little is known about tool-use behaviours that humans can invent individually, i.e. without cultural knowledge. We presented 2- to 3.5-year-old children with 12 problem-solving tasks based on tool-use behaviours shown by great apes. Spontaneous tool use was observed in 11 tasks. Additionally, tasks which occurred more frequently in wild great apes were also solved more frequently by human children. Our results demonstrate great similarity in the spontaneous tool-use abilities of human children and great apes, indicating that the physical cognition underlying tool use shows large overlaps across the great ape species. This suggests that humans are neither born with special physical cognition skills, nor that these skills have degraded due to our species' long reliance of social learning in the tool-use domain.

Western scrub-jays (Aphelocoma californica) solve multiple-string problems by the spatial relation of string and reward.

String-pulling is a widely used paradigm in animal cognition research to assess what animals understand about the functionality of strings as a means to obtain an out-of-reach reward. This study aimed to systematically investigate what rules Western scrub-jays (Aphelocoma californica) use to solve different patterned string tasks, i.e. tasks in which subjects have to choose between two or more strings of which only one is connected to the reward, or where one is more efficient. Arranging strings in a parallel configuration showed that the jays were generally capable of solving multiple-string tasks and acted in a goal-directed manner. The slanted and crossed configurations revealed a reliance on a "proximity rule", that is, a tendency to choose the string-end closest to the reward. When confronted with strings of different lengths attached to rewards at different distances the birds chose according to the reward distance, preferring the reward closest to them, and were sensitive to the movement of the reward, but did not consistently prefer the shorter and therefore more efficient string. Generally, the scrub-jays were successful in tasks where the reward was closest to the string-ends they needed to pull or when string length and reward distance correlated, but the birds had problems when the wrong string-end was closest to the reward or when the food items were in close proximity to each other. These results show that scrub-jays had a partial understanding of the physical principles underlying string-pulling but relied on simpler strategies such as the proximity rule to solve the tasks.

Transfer of physical understanding in a non-tool-using parrot.

Physical cognition has generally been assessed in tool-using species that possess a relatively large brain size, such as corvids and apes. Parrots, like corvids and apes, also have large relative brain sizes, yet although parrots rarely use tools in the wild, growing evidence suggests comparable performances on physical cognition tasks. It is, however, unclear whether success on such tasks is facilitated by previous experience and training procedures. We therefore investigated physical comprehension of object relationships in two non-tool-using species of captive neotropical parrots on a new means-end paradigm, the Trap-Gaps task, using unfamiliar materials and modified training procedures that precluded procedural cues. Red-shouldered macaws (Diopsittaca nobilis) and black-headed caiques (Pionites melanocephala) were presented with an initial task that required them to discriminate between pulling food trays through gaps while attending to the respective width of the gaps and size of the trays. Subjects were then presented with a novel, but functionally equivalent, transfer task. Six of eight birds solved the initial task through trial-and-error learning. Four of these six birds solved the transfer task, with one caique demonstrating spontaneous comprehension. These findings suggest that non-tool-using parrots may possess capacities for sophisticated physical cognition by generalising previously learned rules across novel problems.

A novel form of spontaneous tool use displayed by several captive greater vasa parrots (Coracopsis vasa).

Parrots are frequently cited for their sophisticated problem-solving abilities, but cases of habitual tool use among psittacines are scarce. We report the first evidence, to our knowledge, of tool use by greater vasa parrots (Coracopsis vasa). Several members of a captive population spontaneously adopted a novel tool-using technique by using pebbles and date pits either (i) to scrape on the inner surface of seashells, subsequently licking the resulting calcium powder from the tool, or (ii) as a wedge to break off smaller pieces of the shell for ingestion. Tool use occurred most frequently just prior to the breeding season, during which time numerous instances of tool transfer were also documented. These observations provide new insights into the tool-using capabilities of parrots and highlight the greater vasa parrot as a species of interest for studies of physical cognition.

Development of planning in 4- to 10-year-old children: reducing inhibitory demands does not improve performance.

Currently, there are relatively few tasks suitable for testing planned problem solving in children. We presented 4- to 10-year-old children (N=172) with two planning tasks (sequential planning and advance planning) using the paddle-box apparatus, which was originally designed to investigate the planning skills of nonhuman apes. First, we were interested in the development of children's performance in the two tasks and whether the strategies children used to succeed differed among age groups. Performance improved significantly across age groups in both tasks. Strategies for success in the advance planning task differed among age groups, with 4- and 5-year-olds performing more excess actions, and a greater proportion of irrelevant excess actions, than older children. Findings are discussed in relation to the development of performance in tower tasks, which are a commonly used test of planning ability in humans. Second, based on previous findings with apes, we predicted that introducing measures to reduce the inhibitory demands of the advance planning task would improve children's performance. Therefore, in this study we introduced two methodological alterations that have been shown to improve children's performance in other tasks with inhibitory demands: (a) imposing a short delay before a child is allowed to act and (b) replacing reward items with symbolic tokens. Surprisingly, neither of these measures improved the performance of children in any of the age groups, suggesting that, contrary to our prediction, inhibitory control might not be a key performance-limiting factor in the advance planning paddle-box task.

The role of action concepts in physical reasoning: insights from late childhood.

A fundamental component of human cognition is the ability to intuitively reason about behaviours of objects and systems in the physical world without resorting to explicit scientific knowledge. This skill was traditionally considered a symbolic process. However, in the last decades, there has been a shift towards ideas of embodiment, suggesting that accessing physical knowledge and predicting physical outcomes is grounded in bodily interactions with the environment. Infants and children, who learn mainly through their embodied experiences, serve as a model to probe the link between reasoning and physical concepts. Here, we tested school-aged children (5- to 15-year-olds) in online reasoning games that involve different physical action concepts such as supporting, launching and clearing. We assessed changes in children's performance and strategies over development and their relationships with the different action concepts. Children reasoned more accurately in problems that involved supporting actions compared to launching or clearing actions. Moreover, when children failed, they were more strategic in subsequent attempts when problems involved support rather than launching or clearing. Children improved with age, but improvements differed across action concepts. Our findings suggest that accessing physical knowledge and predicting physical events are affected by action concepts, and those effects change over development. This article is part of the theme issue 'Minds in movement: embodied cognition in the age of artificial intelligence'.

Chimpanzees' (Pan troglodytes) problem-solving skills are influenced by housing facility and captive care duration.

Although a large body of primate cognition research is done in captive institutions, little is known about how much individuals from different facilities vary in their experiences and cognitive skills. Here we present the results of an experimental study investigating how physical cognitive skills vary between chimpanzees in relation to captive settings and their time in captivity. We tested 59 chimpanzees housed at two different captive facilities (a rehabilitation center (sanctuary) and a zoo) in three problem-solving tasks. Our results showed that chimpanzees at the two housing facilities significantly differed in overall task performance. On average, the sanctuary chimpanzees outperformed the chimpanzees housed at the zoo in the detour reaching task and the honey trap task. However, the zoo chimpanzees performed slightly better on average in the learning task. We propose that, for this particular sample, the documented differences result from a combination of factors, such as prior experience with cognitive testing, motivation levels and varying degrees of human exposure. Within the sanctuary sample, we found that chimpanzees who arrived at an earlier age at the sanctuary and had therefore spent a larger percentage of their lives in a captive environment, were better problem-solvers than those that arrived at a later age to the sanctuary. Thus, rehabilitation and time in captivity contributed to improved physical cognitive skills in sanctuary chimpanzees. Our results highlight the importance of studying intraspecific variation and the effect that previous experience and living conditions might have on physical cognitive skills in non-human apes. Accordingly, we should be cautious when extrapolating findings of cognitive studies from one population to the species as a whole.