Animal Cognition

Resource handling time is an important cost to foraging animals and this is especially true for low-ranking individuals who may be displaced before they acquire rewards. Handling may also require visual attention that can lead to a trade-off between scanning the environment and focusing on food acquisition. We ran a foraging experiment on wild vervet monkeys where we simultaneously offered them three puzzle boxes in a straight-line array that increased sequentially in handling difficulty and reward value. We found that differences in competitive ability had a large impact on access to long-handling time resources and willingness to invest time in costly problem-solving when social interference was possible. Resources with lower handling time and smaller rewards were accessed by individuals of all dominance ranks but only dominant monkeys that did not fear displacement attempted the high handling time puzzle box. Handling time improved with experience at each puzzle box and contrary to predictions, when vigilance rates were higher, monkeys were also faster at solving puzzle boxes. Thus, there is not necessarily a trade-off between attention to foraging tasks that require handling and vigilance, if a high scanning frequency to both can be balanced with limited scanning duration. These results have important implications for understanding the decision-making of foraging animals that live in competitive groups and the diets of individuals, which may vary with dominance when important resources require long handling times.

Animals often display active decision-making while navigating noisy habitats, for varied purposes like foraging, finding shelter, evading predators or finding suitable mates. While some responses are hardwired, following fixed action patterns, others are more flexible, and display context-dependent plasticity. Animals might weigh behavioural consistency against adaptive plasticity when facing recurrent ecological difficulties, but the nuanced mechanisms orchestrating this balance is not well understood. We investigated this question by repeatedly exposing 47 free-ranging dogs to chicken contaminated with lemon solution, an aversive but edible and familiar food source, across four consecutive trials. By integrating behavioural frequencies, latency measures, feeding efficiency, interaction-time and location effects, repeatability analyses, and behavioural sequence structure, we examined how repeated experience influenced foraging behaviour. Contrary to expectations of broad behavioural reorganisation, most behaviours remained stable. Initial approach latency was strongly repeatable within individuals with no detectable influence of sex or trial number across the population. This suggests that how an animal first assesses risk is primarily shaped by stable personality traits rather than momentary external factors. Eating behaviour and bowl nudging also changed selectively across exposures, with modest sex differences in several behaviours. Behavioural sequences became increasingly similar over time, while several behaviours retained consistent individual differences, indicating persistent behavioural tendencies. Our results collectively demonstrated that repeated aversive exposure selectively modified a few critical decision points rather than the entire behavioural repertoire. This implies that animals can adapt by making small adjustments to specific behaviours while maintaining the stability of their overall behavioural architectures. Animals seem to assess risk separately from how well they use resources. That separation may help explain how stable personalities can coexist with the fast, flexible decisions animals need in unpredictable environments shaped by humans.

Human-influenced environments pose challenges but also provide wildlife with anthropogenic resources. Individuals vary widely in their ability to exploit such resources, often as a function of behavioural type. However, we lack a clear understanding of how variation in behavioural traits influences stages of resource exploitation required to use anthropogenic resources. Using fully automated foraging puzzles, we examined how boldness and sociability influenced three aspects of resource exploitation - discovery, problem-solving, and overall performance - in two wild populations of California ground squirrels (Otospermophilus beecheyi). Bolder individuals discovered the resource earlier, solved the task faster and achieved higher performance, indicating that boldness promotes efficient exploitation of anthropogenic resources across multiple stages in the process. Greater sociability and more opportunities to observe conspecifics solving the task led to faster problem-solving, consistent with evidence for observational learning. Squirrels in the recreational-use population - with regular exposure to humans and anthropogenic food - were faster to discover the resource than those in a trail-use population - where human exposure was transient and no anthropogenic food available. Problem-solving latency and performance were consistent between populations. Our findings highlight how individual variation in behavioural traits drives performance in novel ecological contexts, providing a mechanistic understanding of behavioural plasticity in human-influenced environments.

Prey capture has long served as a model of complex, multimodal behavior with ethological relevance. In this context, the ability of laboratory animals to capture fleeing prey has been investigated in terms of the sensory systems and neuroanatomical regions required for successful capture. While the ability to hunt is experience-dependent, prey capture is often assumed to be a reflexive behavior, governed by innate neural circuits and a basic drive to obtain food. Whether hunting performance depends on complex learning and cognitive strategy remains largely untested, in part because it is difficult to systematically manipulate the difficulty of predator-prey interactions. To address this, we varied ambient temperature across thousands of trials in which laboratory mice pursued cold-blooded cockroaches, whose movement speed was directly determined by body temperature. As expected, capture difficulty increased as a function of temperature. To test whether mice rely on external cues to adjust their predatory behavior, we then mismatched the ambient temperature and that of the prey, finding that mice learn to use relevant environmental information to inform their hunting strategy. Our results suggest that animals can extract and apply meaningful information from predictive environmental cues to adapt their learned behavior.

Learning locations by integrating external reference points, or place learning, is a form of navigation strongly associated with the mammalian hippocampus and its homologs in other vertebrates. In contrast, cue learning relies on individual environmental features and can occur independently of the hippocampus. Despite growing interest in amphibian spatial cognition and association of spatial functions with the medial pallium brain region, how amphibian brains support different spatial learning strategies remains unknown. We designed a four-arm maze to test place, cue, and turn-direction learning in the cane toad (Rhinella marina), and paired maze trials with visualization of brain activity through pS6 immunohistochemistry. Toads improved performance over time only in the place task but exhibited consistently faster exit times in the cue task. Place task toads displayed elevated brain activity only in the dorsal and lateral palliums. These observations demonstrate that amphibians can acquire spatial navigation strategies through distinct behavioral paradigms and reveal the diverse pallial contributions underlying vertebrate spatial cognition.

Adapting behaviour when reward contingencies change is a core function of cognitive control, but the underlying trial-by-trial computations are hard to observe when tasks cue each rule or allow only one switch per session. We developed the Feature-Rule Switching Task (FRST), in which common marmosets (Callithrix jacchus) repeatedly switch, without cues and under a fixed stimulus set, among the visual features that earn reward, inferring each switch from feedback alone. All four animals acquired the task within two days and sustained several switches per session across months of testing. A reinforcement-learning model with a learned weighting of stimulus dimensions best explained choices in every animal, outperforming complexity-matched perseveration controls. Learning-rate estimates fell within the human range, and in almost every session the animals weighted a stimulus dimension rather than individual features alone, with a dimensional commitment comparable in strength to that of humans. FRST thus provides a primate paradigm, amenable to laminar recording, for tracking the dynamics of cognitive flexibility.

Animals differ in how they acquire and retain information, and modify their behavior while interacting with their environment. These differences have been linked to personality type, yet how personality type and sex influence reward-associated memory is not well understood. We investigated the effects of personality type, sex, days of conditioning, and food reward on learning and memory retention in zebrafish using the conditioned place preference paradigm. We conditioned zebrafish to associate a visual background with a food reward across eleven sessions and assessed memory recall seven days later by measuring duration, frequency of entry, and latency to first entry in the conditioned zone. We observed that bold fish had early conditioned zone engagement, whereas shy zebrafish gradually increased their behavior into the conditioned zone with repeated conditioning. With food reward, preference for the conditioned zone increased after 7 and 11 days post-conditioning relative to baseline. Seven days after the last conditioning with food reward, the preference for conditioned zone remained unchanged in shy fish. This suggests that shy fish retain the ability to recall the learned association for at least 7 days after conditioning. Overall, personality, food reward, and sex influence behavioral changes in food reward-associated cognition, with shy-treated males showing gradual increases in behavior into the conditioned zone. These findings suggest that personality and sex differences may depend on the nature and valence of the associative task to recall. Thus, this study emphasizes the importance of assessing memory retention alongside learning acquisition to understand the relationship between personality types and cognition.

Decision-making is key to survival, with choices and behaviour typically shaped by evolutionary pressures to enhance fitness and minimize loss. Inequity aversion, the tendency to respond negatively to unequal reward distributions, is therefore unexpected, as acts of fairness may be costly to the individual in the short term. In social, cooperative species, however, fairness may enhance long-term benefits through stable social interactions. Inequity aversion is believed to promote cooperative social structures and has only been demonstrated in certain social species. Yet, solitary species have not been tested, and whether this behaviour depends on social cooperative structures is unknown. Examining solitary species could clarify whether inequity aversion reflects an adaptation to cooperation or a general cognitive capacity linked to social comparison and reward evaluation. We studied inequity aversion in polar bears (Ursus maritimus), a largely solitary species, using three complementary, non-invasive cognition tests on seven bears from two zoos in the Netherlands. (1) An impunity experiment assessing effort-based inequity, in which bears performed a standardized action to obtain food rewards. (2) A choice-based experiment testing active selection between equal and unequal reward distributions. (3) A no-task control experiment assessing responses to unequal reward distribution without effort. Under effort-based inequity, bears reduced task performance and increased task latency when a partner received a superior reward. Unequal reward distributions did not affect reward acceptance independent of effort. In the choice-based paradigm, bears preferentially selected equal reward distributions with a partner present. Although generalizability to wild populations is limited, these findings demonstrate behavioural aversion of social inequity in a non-cooperative species. Inequality aversion may therefore not be restricted to social, cooperative animals but instead reflect a broadly distributed cognitive mechanism that emerges under relevant social conditions. These findings challenge current theories and advance our understanding of the cognitive foundations of fairness-related behaviour in animals.

Foraging decisions often depend on net energy intake, but the need for specific micronutrients means that foods are not always comparable based on energy content. Although sodium is essential for many physiological functions, many organisms face challenges obtaining it, especially those relying on a plant-based diet. Flowering plants often lure pollinators by manipulating nectar carbohydrate content, but nectar sodium might also influence pollinator choice. We experimentally tested how sodium addition and nectar carbohydrate concentration interact to affect bumble bee foraging decisions by allowing bees to forage on artificial flowers containing either low- or high-sucrose nectar, with or without sodium. We also explored whether sugar and sodium content in floral nectar trade-off across plant species. Bees preferred sodium-enriched nectar and preference nearly doubled when sucrose concentration was low. Furthermore, bees switched more often from flowers without sodium to those with sodium when sucrose concentration was low. Given increased fidelity for artificial flowers with nectar sodium, sodium may enhance plant fitness by increasing conspecific pollen receipt. Yet despite behavioral evidence that sodium enhances the perceived quality of low-sucrose nectar for bees, we found no evidence for a nectar sugar-sodium trade-off across plant species. We discuss implications for plant-pollinator interactions and flower evolution.

Exploration is fundamental to how children learn about the world. To explore a new environment comprehensively, a child must navigate without getting lost while simultaneously attending to salient information along the way. Both abilities improve across childhood, yet most studies examine them in isolation, often in constrained spaces with limited sensory complexity. Here, we asked 7- to 14-year-old children (N=42) and young adults (18-22; N=42) to freely explore a large furniture store in immersive virtual reality, which contained many objects and attention-capturing multisensory events. Surprisingly, despite traveling comparable distances, children explored less - they covered less spatial area and examined fewer unique objects than adults. Fine-grained analyses of their behaviors showed that the reduction in exploration was due to children spending more time backtracking and children taking longer to adapt to salient multisensory objects, which were uncorrelated across individuals after controlling for age and independently predicted exploration. However, only backtracking duration predicted success in a subsequent search task, demonstrating a link between navigational success during exploration and the quality of learned spatial representations for later use. Together, the results demonstrate that children explore less comprehensively than adults in large-scale spaces because they are susceptible to getting lost and perseverating on salient events nested within a large environment, but that these behaviors change with age. This suggests that while children are widely considered to be curious explorers within local environments, their ability to do so within larger environments is shaped by the maturation of cognitive systems that integrate experiences over complex contexts.

Mice are widely used to study the neural basis of perceptual decision-making, often with the ultimate goal of extrapolating insights to humans. Yet, whether humans and mice rely on similar computational strategies remains rarely tested, partly because the two species acquire knowledge differently. Animals learn from experience, while humans usually receive verbal task instructions. Here, we explicitly contrast these modes of learning in humans who perform a perceptual decision-making task adapted from a standardized rodent task. Specifically, we present an open behavioral dataset from 80 human participants who performed 600 trials of a visual contrast-discrimination task, matched to the standardized International Brain Laboratory task for mice (International Brain Laboratory et al., 2021, 2025). Participants either received verbal instructions ('Instructed') or were only told to discover the rules of the task by playing ('Uninstructed'). The dataset contains choices, response times, continuous cursor trajectories, eye-tracking and pupil recordings. Descriptive analyses show that Uninstructed participants were less accurate and had higher lapse rates than Instructed participants. They improved over the first 200 trials, approaching the performance of Instructed participants. These data provide a benchmark for testing whether computational strategies identified in mice also describe human behavior.

In vertebrates, top-down visual attention is a cognitive process where internal goals modulate the tuning of peripheral sensory systems. This leads to increased perceived contrast to both goal-relevant objects and areas of the visual field that are attended. Such a system would also be beneficial to bees, enabling them to detect and recognise the most profitable flowers in their environment. We tested whether bumblebees possess a top-down attentional system resembling that seen in vertebrates. We trained two groups of bees to collect rewards under high contrast targets. To potentially induce a difference in attention while searching for the targets, one group received a higher concentration of sucrose rewards compared to the other. During tests, the targets were presented with a series of lower contrasts to measure the contrast sensitivity curves of the bees induced by the different learnt reward levels. We predicted a stronger effect of any attention-like process on contrast sensitivity in the high reward group. We also repeated this experiment with the neonicotinoid pesticide imidacloprid dissolved in the sucrose rewards to test whether this affects bee attention. Across all test contrasts, higher rewards significantly increased bee accuracy when locating targets, lowered contrast thresholds and reduced the latency to make first choices. Imidacloprid reduced bee accuracy but did not influence first choice latency. These results suggest that learnt floral rewards can influence bee behavioural contrast sensitivity in a manner resembling vertebrate top-down attention and that imidacloprid may modulate this through effects on their nervous system.

When food resources provide reward while simultaneously imposing risk, animals can regulate their behaviour to continue exploiting them while reducing associated costs. Angling presents such a feeding situation, in which obtaining bait can result in hooking and capture. Although fish become less vulnerable to angling following capture experience, the behavioural changes underlying this reduced vulnerability remain poorly understood. Here, we used fine-scale video observations to examine feeding behaviour of red sea bream (Pagrus major) toward angling gear before and after capture experience. Naive fish rapidly swallowed the bait and were subsequently captured, whereas capture-experienced fish continued to approach and feed on the gear but shifted from swallowing to biting. This feeding-mode shift may allow fish to maintain exploitation of the bait while reducing capture risk. Our observations provide a behavioural explanation for reduced angling vulnerability and suggest that direct experience with capture risk can alter how fish exploit a rewarding food resource.

Reinforcement learning theory formulates distinct decision-making strategies, including reactive model-free and deliberative model-based strategies. This study investigates how mice adjust their reinforcement learning strategies while learning decision-making in dynamic environments. Unlike previous studies that focused on behaviors after extensive training periods, we analyzed changes in learning strategies in the course of training of a two-step decision-making task with probabilistic state transition and fluctuating reward probabilities. Our statistical behavioral analysis showed that the stay-probability following common and rare transitions diverged with training, a signature of strategies that utilize knowledge of task structure. We fit various reinforcement learning strategies to behavioral data and found that structure-informed strategies became increasingly dominant in their behaviors during training. Whereas previous studies emphasized transition from goal-directed to habitual strategies after extensive training, which were often associated with model-based and model-free strategies, respectively, our results newly demonstrate a shift from model-free to structure-informed strategies in early training in mice. Author summaryReinforcement learning theory allows us to examine how we make decisions and what approaches we use to optimize rewards. Most previous research, however, has examined animal behavior only after extensive training. Here we analyzed how mice adjust their reinforcement learning strategies as they are trained in a two-step decision-making task. Initially, mice relied on reactive model-free strategies, but as training progressed, their behavior began to incorporate knowledge of task structure. While previous studies suggested transition from model-based to model-free strategies with extensive training, our study revealed the opposite in the early stage of training.

Environmental predictability influences the value of information acquired through experience, yet relatively little is known about how instability in resource characteristics influences behavioural organisation during foraging. We tested whether repeated changes in floral orientation, a manipulation of environmental predictability, affect pollen foraging in bumblebees (Bombus terrestris) by exposing naive workers to either stable floral conditions (single flower orientation) or repeated inter-trial changes in flower orientation (three alternating flower orientations), under constant resource availability. We quantified pollen collection, foraging efficiency, revisitation behaviour, floral coverage and sonication behaviour across three successive foraging trials of either constant or variable flower orientation, before assessing performance in a common post-treatment preference test in which bees were offered all three flower orientations and higher pollen rewards. Environmental instability altered the organisation of foraging behaviour. Bees exposed to unstable floral conditions progressively reduced flower revisitation behaviour and were less likely to perform sonication, although floral coverage, defined as the number of unique flowers visited, remained unchanged. Contrary to our predictions, instability had only weak immediate effects on pollen acquisition and foraging efficiency compared to bees tested under stable conditions. However, previous exposure to instability generated carry-over effects in the common post-treatment preference test. Bees previously exposed to unstable conditions were significantly less likely to return with pollen and consequently collected less pollen overall than bees previously exposed to stable conditions. Our results demonstrate that environmental instability can influence pollen foraging in ways that are not captured by immediate measures of performance. Although behavioural adjustments appeared to buffer short-term consequences during repeated foraging trials, carry-over effects were evident when bees were later tested in a common high-reward, multi-orientation floral array. These findings highlight the importance of considering both behavioural flexibility and carry-over effects when evaluating how organisms respond to changing environments.

Recent research has suggested that the availability of non-instrumental information about the outcome of a risky choice increases risk appetite. In this study, we aimed to perform a conceptual replication of these findings and to examine the cognitive mechanisms underlying this effect. Across two experiments (N = 150, 102), we presented participants with mathematically fair gambles and allowed them to choose the size of their bet. Between trials, we varied the presence of non-instrumental information that would reveal the outcome ahead of time. In both experiments, we did not find consistent evidence for an effect of the availability of non-instrumental information on bet size. These findings suggest that the previously reported effects of non-instrumental information on risk appetite may have been an idiosyncratic feature of experimental design, rather than a more general phenomenon that characterises human decision making under risk.

Defensive survival behaviors are shaped by both danger attributes and internal physiological states. Well-characterized defensive responses depend not only on whether danger is potential or perceived, but also on its spatial and or temporal proximity. The relationship between danger imminence and defensive behaviors is referred to as the threat imminence continuum, which provides a conceptual framework bridging ecology and behavioral neuroscience. Although this theoretical framework is gaining momentum, another much less explored danger attribute is the extent to which the environment may be controllable. Most paradigms assessing defensive behaviors rely on conditioned stimuli as danger warning-signals in stable environments that hamper the study of environment controllability and its impact on defensive behaviors. To address this question, we repurposed the active place avoidance (APA) paradigm combined with supervised machine learning to characterize multivariate exploratory and defensive behaviors in male and female mice. One important benefit of APA is that it allows the study of active avoidance when the platform is rotating as well as deliberative avoidance when the platform is immobile. As such, APA is very well suited to test the impact of environment controllability on the expression of defensive behaviors. Using this unified preparation, we systematically compared quantitative danger attributes (punishment intensity, behavioral agency) across danger states (absence, presence and omission of punishment) and learning experience (early and late learning stages), while considering sex as a biological variable. Our results demonstrate that defensive behaviors depend on punishment intensity in a sex-dependent manner in APA. Defensive responses can be classified as proactive (risk assessment) or reactive (freezing, escape) in APA. Their response probability depends on behavioral agency, is mutually exclusive and depends on danger spatial proximity. Acute treatment with norepinephrine transporter blocker selectively increases reactive defensive behaviors while sparing proactive defensive behavior. Collectively, these results provide experimental evidence that environmental stability calibrate proactive and reactive defensive behaviors whose neural correlates may be divergent.

Objective and efficient methods for investigating animal's mental states are important for advancing animal welfare research. Behavioural observations and physiological measures provide valuable information but may not fully capture affective states. Judgment Bias Tasks (JBTs) offer a promising method to infer affective states from animals' responses to ambiguous stimuli, as distressed individuals show more pessimistic tendencies. However, the application of JBTs outside research settings may be limited due to time-consuming training and methodological or infrastructural requirements. With the present study, we aimed to test the feasibility of a simple and accessible JBT for non-human primates and to investigate its sensitivity to affective states. Eight brown capuchins (Sapajus apella) and eleven Tonkean macaques (Macaca tonkeana) were tested under positive and negative affective conditions. Individuals were trained to discriminate between never rewarded conditioned stimuli and always rewarded ones. They were then presented with ambiguous probes they had never seen before to analyse their responses to ambiguity. The discrimination training required, on average, about 12 sessions lasting a few minutes each to successfully train every individual animal. Both species significantly discriminated between conditioned stimuli and ambiguous probes. Additionally, macaques showed a greater tendency to interact with the never rewarded condition stimuli and the probes under positive conditions during early sessions. In capuchins, however, the affective conditions presented had no significant effect, although the descriptive analysis tended to show that the latency to investigate the probes was greater in the negative condition. This study also highlights key methodological considerations, such as a decrease in ambiguity across sessions. Overall, the present results suggest that, with a few improvements, this method could be effectively implemented to assess affective states of primates under human care, supporting this tool as a valuable candidate for welfare assessment.

Shorebirds provide an excellent model for investigating the relationship between bill morphology and food acquisition. Food acquisition comprises three successive behavioural stages: foraging (locomotion and prey capture), feeding (food handling and transport), and swallowing. During food transport, these birds use two non-lingual mechanisms, surface-tension transport (ST) and ballistic transport (BT), whose characteristics depends on the kinematics of head and beak movements and the physical properties of the food. We investigated the behavioural flexibility of these transport mechanisms in captive and free-ranging individuals of two species with contrasting beak morphologies: the Pied avocet (Recurvirostra avosetta) and the Black-winged stilt (Himantopus himantopus). Although these species have beaks of a similar size, avocets are distinguished by their upward-curved beaks, whereas stilts have a rather straight beak. We examined the effects of food water content and the presence or absence of water in the beak on food transport by quantifying maximum gape, maximum head displacement, and maximum head velocity. Transport kinematics were jointly influenced by food properties and water availability in the beak. Moist food improved transport performance in both species, whereas dry food required compensatory increases in gape amplitude and head movements, demonstrating that neither ST nor BT constitutes a fixed behavioural sequence. Species also differed consistently in their transport strategies: Black-winged stilts relied on slower, larger-amplitude head movements, whereas Pied Avocets exhibited faster, more precise movements, particularly when water was present. These findings demonstrate that ST and BT share common biomechanical foundations while being governed by rapid kinematic adjustments to changing environmental conditions that probably correspond to flexible motor control. This behavioural flexibility in food transport is therefore likely to enhance feeding performance and ecological resilience in the heterogeneous habitats occupied by shorebirds, suggesting that context-dependent modulation of transport behaviour represents an important adaptive feature of these shorebirds.

Understanding the evolution of cognition is fundamental to understanding the origins and nature of our own cognitive abilities, with cross-species comparisons providing a cornerstone of this endeavor. Yet, comparative cognition so far has largely focused on endothermic vertebrates, particularly mammals and birds, while ectotherms--representing a substantial proportion of vertebrate diversity--remain relatively understudied. Moreover, variation in experimental methods makes existing data difficult to compare across species. Big-team science offers a way to overcome these limitations by enabling standardized cognitive testing across broad taxonomic scales. Here, we apply this approach to fishes, using inhibitory control as a proof of concept for standardized comparative testing in 444 individuals across 22 teleost species spanning 19 genera. We demonstrate that standardized cognitive testing across such diversity is feasible, while also revealing substantial heterogeneity in the sample. Performance varied markedly among species, but, strikingly, showed only a weak phylogenetic signal, contrasting with patterns previously reported in mammals and birds. In addition, we detected great within-species variation, including differences across research groups, suggesting that performance may reflect not only species-level evolutionary history but also ontogenetic experience and the capacity to adjust decision rules to local conditions. Together, our findings highlight the importance of both evolutionary and environmental sources of cognitive variation and demonstrate that extending standardized comparative approaches beyond traditionally studied endotherms can reveal a more dynamic and complex picture of cognitive evolution across the vertebrate lineage.