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Non-instructed Motor Skill Learning in Monkeys: Insights from Deep Reinforcement Learning Models

In the field of motor learning, few studies have addressed the case of non-instructed movement sequences learning, as they require long periods of training and data acquisition, and are complex to interpret. In contrast, such problems are readily addressed in machine learning, using artificial agents in simulated environments. To understand the mechanisms that drive the learning behavior of two macaque monkeys in a free-moving multi-target reaching task, we created two Reinforcement Learning (RL) models with different penalty criteria: "Time" reflecting the time spent to perfom a trial, and "Power" integrating the energy cost. The initial phase of the learning process is characterized by a rapid improvement in motor performance for both the 2 monkeys and the 2 models, with hand trajectories becoming shorter and smoother while the velocity gradually increases along trials and sessions. This improvement in motor performance with training is associated with a simplification in the trajectory of the movements performed to achieve the task goal. The monkeys and models show a convergent evolution towards an optimal circular motor path, almost exclusively in counter-clockwise direction, and a persistent inter-trial variability. All these elements contribute to interpreting monkeys learning in the terms of a progressive updating of action-selection patterns, following a classic value iteration scheme as in reinforcement learning. However, in contrast with our models, the monkeys also show a specific variability in the choice of the motor sequences to carry out across trials. This variability reflects a form of "path selection", that is absent in the models. Furthermore, comparing models and behavioral data also reveal sub-optimality in the way monkeys manage the trade-off between optimizing movement duration ("Time") and minimizing its metabolic cost ("Power"), with a tendency to overemphasize one criterion at the detriment of the other one. Overall, this study reveals the subtle interplay between cognitive factors, biomechanical constraints, task achievement and motor efficacy management in motor learning, and highlights the relevance of modeling approaches in revealing the respective contribution of the different elements at play. Author summaryThe way in which animals and humans learn new motor skills through free exploratory movements sequences solely governed by success or failure outcomes is not yet fully understood. Recent advances in machine learning techniques for continuous action spaces led us to construct a motor learning model investigate how animals progressively enhance the efficiency of their behaviors through numerous trials and errors. This study conducts a comprehensive comparison between deep learning models and experimental data from monkey behavior. Notably, we show that the progressive refinement of motor sequences, as they are observed in the animals, do not require the implementation of a complete model of their environment. Rather, it merely requires the capacity to anticipate both movement costs and final reward a few steps ahead in the future following a value iteration principle. Furthermore, the systematic deviations exhibited by the monkeys with respect to the computational model inform us on the presence of individual preferences in either minimizing the duration or the energy consumption, and also on the involvement of alternative "cognitive" strategies.

animal behavior and cognition↗

Bumblebee queens differ in brain morphology but not learning performance across life stages

Animal cognition and brain morphology can vary between individuals and across a lifetime as a function of social and ecological requirements. Bumblebees have distinct ecological pressures acting upon individuals at different times: young queens (gynes) and workers share similar challenges as they both forage as part of the colony, but only queens overwinter and start a colony the following season, switching to a reproductive stage. Here we compared bumblebee (Bombus impatiens) visual learning and brain morphology across age-matched workers and gynes and older reproductive queens. We expected foraging-stage gynes to be better at visual learning than reproductive queens and visual regions to be reduced in the reproductive queens, in line with work in other social insects. However, we found that gynes and queens performed similarly, while both performed better than workers. We also found that reproductive queens had larger olfactory regions (antennal lobes) than gynes, while visual regions (medulla and lobula) did not differ, indicating a greater investment in olfactory regions in this later reproductive stage. Brain regions also scaled differently with body size for gynes and workers. Overall, our results provide behavioral and morphological evidence that social and ecological roles in a colony shape cognition and neural investment.

animal behavior and cognition↗

What's in it for the dogs? Assessing the outcomes of a prison-based dog training program from an animal behavior and welfare perspective

Prison-Based Dog Training Programs (PBDTPs) are gaining popularity across the world for their benefits to inmates in terms of mental and emotional health, and reduced recidivism. However, research on the implications for the dogs is limited. The aim of the present study was to assess the outcomes of a PBDTP from an animal behavior and welfare perspective. Shelter dogs (N=42) were transported to and from the prisons twice a week where they received training from inmates for a total duration of 12 weeks. Dogs were tested for potential improvements in socialization and handling skills and basic training skills using standard tests (Temperament Test and Basic Education Test). Dog welfare was assessed using behavioral, physiological, and cognitive measures, namely stress-related behaviors and overall behavior states during training sessions, salivary cortisol levels, and cognitive bias tests. Results showed that participating in the PBDTP improved the dogs sociability towards humans and increased playful behavior, and improved basic training skills including not jumping on people, walking on a leash without pulling, responding to commands (sit, down, and staying in place) and staying calm when separated from the handler. Furthermore, behaviors indicative of stress were generally rare during training sessions and no impact of the PBDTP was found on the levels of salivary cortisol nor on the dogs affective states (as measured with cognitive bias tests). In conclusion, the present study suggests that PBDPTs are beneficial for dogs, with animals showing no indicators of compromised welfare while displaying improved behavior skills, which will likely facilitate their future rehoming.

animal behavior and cognition↗

Capturing Rapid Behavioral Adaptation in an Updated Theory of the Hippocampal Cognitive Map and its Neural Implementation

Animals sometimes adapt their behavior after a single exposure to novel information. Here, we integrate abstract theory with recent mechanistic insights to understand this and other examples of rapid adaptation. We ground our investigation in studies of animal learning in spatial contexts, where it is widely held that learning depends on the formation of a cognitive map reflected in learned representations in hippocampal areas CA1 and CA3. We extend the successor representation (SR) theory of the cognitive map, incorporating the weighting of this representation by perceived salience (PS) signals, and using it to represent future availability of environmental features that allow behavior to be guided flexibly by an animals goals and needs at later times. To capture rapid adaptation, we rely on behavioral time-scale synaptic plasticity (BTSP), a form of plasticity that captures one-shot formation of new place fields. Through simulations, we show that our hippocampal model network can learn the PS-weighted SR rapidly and set up the ability to replay experiences in the environment after a single exposure through BTSP. Spontaneous replay activity during subsequent offline periods strengthens the PS-weighted SR. Predictions of environmental features also develop rapidly, and behavioral simulations show how these learned representations give rise to need-dependent choice, rapid behavioral adaptation to changes in the availability of reward, and one-trial shock avoidance learning.

neuroscience↗

Human infants’ understanding of social imitation: Inferences of affiliation from third party observations

Imitation is ubiquitous in positive social interactions. For adult and child observers, it also supports inferences about the participants in such interactions and their social relationships, but the origins of these inferences are obscure. Do infants attach social significance to this form of interaction? Here we test 4- to 5.5-month-old infants interpretation of imitation, asking if the imitative interactions they observe support inferences of social affiliation across 10 experimental conditions that varied the modality of the imitation (movement vs. sound), the roles of specific characters (imitators vs. targets), the number of characters in the displays (3 vs. 5), and the number of parties initiating affiliative test events (1 vs. 2). These experiments, together with one experiment conducted with 12-month-old infants, yielded three main findings. First, infants expect that characters who engaged in imitation will approach and affiliate with the characters whom they imitated. Second, infants show no evidence of expecting that characters who were targets of imitation will approach and affiliate with their imitators. Third, analyzing imitative interactions is difficult for young infants, whose expectations vary in strength depending on the number of characters to be tracked and the number of affiliative actors to be compared. These findings have implications for our understanding of social imitation, and they provide methods for advancing understanding of other aspects of early social cognitive development.

Animal Behavior and Cognition↗

Learning for angling: an advanced learning capability for avoidance of angling gear in red sea bream juveniles

Angling has been the cause of mortality for fish since ancient. The avoidance learning for angling gear could be considered as a survival strategy against the mortality by angling. Whereas some studies indicated the possibility of avoidance learning for angling gear, most studies investigated the avoidance learning by using groups of fish, in which it is difficult to reveal the process and mechanisms of the learning. The present study elucidated the avoidance learning for angling gear by experiment of single fish in a tank using red sea bream Pagrus major juveniles. Individuals with only once or twice of experience for angling avoided angling gear while showing the feeding motivation for pellets, representing avoidance learning for the angling gear. Most of the experienced individuals avoided the krill attached with a fishing line, but not krill and pellets near the angling gear. Feeding rate for prey on a fishing line at two month after the angling trial demonstrated that approximately half of fish kept the memory for angling gear. A series of experiment for angling gear elucidated that red sea bream juveniles are equipped with considerable learning capability for angling gear, suggesting a cognitive evolution for angling.

animal behavior and cognition↗

Psychophysiological stress induced neurobehavioral changes in adolescent rat and their impact in Adult rat enhanced by cinnamaldehyde

Adolescent is a tricky developmental period were the Physical, cognitive and social changes takes place. Adolescent brain undergoes vigorous maturation and more susceptible to stressors, distresses in the brain development leading to more susceptibility for stress related psychophysiological disorders in later life. The present study explores the association of adolescent stress on same period and on adult. Adolescent rats were restraint to induce psychophysiological stress from PND28 to PND42.Cinnamaldehyde (CA) 25 mg/kg body weight is treated along with the stress. Restraint stress showed more deleterious effect on learning, memory, anxiety, depression and locomotor activity. The result of this study reveal that adolescent stress has impact on that period and even long-lasting impact. But CA may act as antistressor and cognitive enhancer.

animal behavior and cognition↗

Pessimistic and optimistic cognitive biases in cockroaches

Understanding the subjective experience of animals is key for how we manage them and for mapping the evolution of cognitive complexity. Emotional and affective are fundamental to these goals as their presence indicates the potential for subjective experiences and suffering. However, until recently the potential for affective states in invertebrates has been underappreciated, especially for positive affective states where research is limited to a single order. Here, we adapt cognitive bias tests to examine both negative (pessimistic) and positive (optimistic) bias in cockroaches (Blaptica dubia) in order to expand the taxonomic reach of affective state research and to provide a rare test for positive affective states in an invertebrate. We trained cockroaches to associate two different odours with either positive or negative reinforcement and then assessed their reactions to ambiguous stimuli (a range of mixtures of the trained odours). We found that cockroaches exposed to light were more pessimistic (negative affective state) than controls, while those exposed to the scent of opposite sex conspecifics were marginally more optimistic (positive affective state) than controls. We therefore demonstrate the potential for affective states in cockroaches and for positive affective states in invertebrates more widely, indicating a hereto unexpected level of cognitive complexity.

animal behavior and cognition↗

Revisiting the social brain hypothesis: contest duration depends on loser’s brain size

BackgroundBrain size is expected to evolve by a balance between cognitive benefits and energetic costs. Several influential hypotheses have suggested that large brains may be especially beneficial in social contexts. Group living and competition may pose unique cognitive challenges to individuals and favor the evolution of increased cognitive ability. Evidence comes from comparative studies on the link between social complexity and brain morphology, but the strength of empirical support has recently been challenged. In addition, the behavioral mechanisms that would link cognitive ability to sociality are rarely studied. Here we take an alternative approach and investigate experimentally how brain size can relate to the social competence of individuals within species, a problem that so far has remained unresolved. We use the unique guppy brain size selection line model system to evaluate whether large brains are advantageous by allowing individuals to better assess their performance in a social contest situation. Based on theoretical literature, we predict that contest duration should depend on the brain size of the loser, as it is the capitulation of the losing individual that ends the fight.\n\nResultsFirst, we show that studying the movement of competitors during contests allows for precise estimation of the dominance timeline in guppies, even when overt aggression is typically one-sided and delayed. Second, we staged contests between pairs of male that had been artificially selected for large and small relative brain size, with demonstrated differences in cognitive ability. We show that dominance was established much earlier in contests with large-brained losers, whereas the brain size of the winner had no effect. Following our prediction, large-brained individuals gave up more quickly when they were going to lose.\n\nConclusionsThese results suggest that large-brained individuals assess their performance in contests better and that social competence indeed can depend on brain size. Conflict resolution may therefore be an important behavioral mechanism behind macro-evolutionary patterns between sociality and brain size. Since conflict is ubiquitous among group-living animals, the possible effects of the social environment on the evolution of cognition may be more broadly applicable than previously thought.

animal behavior and cognition↗

Reusable modular architecture enables flexible cognitive operations

Complex behaviors are thought to be built by combining simpler cognitive components. Computational modeling has shown that artificial neural networks can perform a variety of tasks by flexibly combining functional modules, each specialized for a specific computation, to construct a complex task. However, it is unknown whether reusable modular networks are found in the brain. Here, we show that mice performing a delayed match-to-sample with delayed report (DMS-dr) task reuse neuronal subspaces that were specialized for stimulus processing and memory maintenance. These subspaces were reused during the task to represent new stimulus inputs and different types of memories, respectively. Clustering analyses showed each subspace was supported by a distinct cluster of neurons in prefrontal cortex and parietal cortex. Studying artificial recurrent networks constrained to neural data found silencing specific clusters disrupted specific computations, consistent with a modular and reusable organization. Altogether, our findings show that the brain can flexibly reuse computational components to perform a complex cognitive task.

animal behavior and cognition↗

Cognitive impairments in a mouse model for Huntington's disease correlate with presymptomatic locomotion and number of CAG repeats

Huntingtons disease (HD) is a progressive neurodegenerative disorder caused by an expanded CAG repeat in the huntingtin (HTT) gene. The disease is characterized by movement disorders, and it also presents with personality changes, including apathy and aggression, along with cognitive decline. While most animal models for HD have been validated for motor deficits, less is known about alterations in other behavioral functions. Here, we performed a longitudinal study to analyze the behavior of a knock-in mouse model of HD with a chimeric mouse/human exon 1 containing 140 CAG repeats inserted in the murine huntingtin gene. We specifically inquired about the onset of cognitive impairments in knock-in mice and whether changes in various behavioral functions such as locomotion, anxiety, and cognition correlate at the individual level. Our data indicate that female and male knock-in mice exhibit reductions in body weight, novelty-induced locomotion, and remote spatial memory retrieval. However, social behavior, working, and short-term memory remain unaffected. Within knock-in mice, lower open-field activity correlated with poorer remote memory performance. Moreover, CAG repeat length negatively correlated with locomotor activity and spatial memory, indicating that greater repeat expansion predicts more severe behavioral impairment. These findings identify early affective changes, followed by selective long-term memory and locomotor deficits, in knock-in mice, supporting this model as a useful platform for studying prodromal HD and repeat-length-dependent disease variability. HighlightsO_LICAG140 knock-in mice show early anxiety, later reduced locomotion and memory deficits C_LIO_LILong-term and remote memory are impaired while short-term and working memory are spared C_LIO_LILower locomotion at the age of 8 months correlates with poorer memory at 14 months of age in individual CAG140 knock-in mice C_LIO_LIGreater CAG repeat length predicts worse locomotion and memory C_LI

animal behavior and cognition↗

Negative memory engrams in the hippocampus enhance the susceptibility to chronic social defeat stress

The hippocampus has been highly implicated in depression symptoms. Recent findings suggest that the expression and susceptibility of depression symptoms are related to the enhanced functioning of the hippocampus. We reasoned that hippocampal engrams, which represent ensembles of neurons with increased activity after memory formation, could underlie some contributions of the hippocampus to depression symptoms. Using the chronic social defeat stress (CSDS) model, we examined social defeat-related hippocampal engrams in mice that are either susceptible or resilient to the stressor. TetTag mice were used to label social defeat-related hippocampal ensembles by LacZ. Engram cells correspond to ensembles that were reactivated by the same stressor.\n\nCompared to resilient and non-stressed control mice, we found that in both the dorsal and ventral hippocampal CA1 regions, susceptible mice exhibited a higher reactivation of social defeat-related LacZ-labeled cells (i.e. engram cells). The density of CA1 engram cells correlated with the level of social avoidance. Using DREADD to reactivate social defeat-related but not neutral contextual stimuli-related CA1 engram cells decreased social interaction. Increased engram cells in susceptible mice were region specific and could not be found in the dentate gyrus. Susceptible mice exhibited more negative stimuli-, but not neutral stimuli-, related CA1 engram cells than resilient mice in the dorsal hippocampus. Finally, chronic, but not a short and subthreshold, social defeat protocol was necessary to increase CA1 engram cell density. Together, our findings reveal that the susceptibility to CSDS is regulated by hippocampal CA1 engrams for negative memory. Hippocampal engrams for negative memory may underlie the vulnerability and expression of cognitive symptoms in depression.\n\nSignificance statementWe provided evidence that negative memory hippocampal engrams contribute to the susceptibility to developing depression-related behavior after chronic social defeat stress. The activation of positive memory engrams have been shown to alleviate depression-related behaviors, while our findings reveal the pathological roles of negative memory engrams that could lead to those behaviors. Increased negative memory engrams could be a downstream effect of the reported high hippocampal activity in animal models and patients with depression. Unlike affective symptoms, we know much less about the cellular mechanisms of the cognitive symptoms of depression. Given the crucial roles of hippocampal engrams in memory formation, enhanced reactivation of negative memory engrams could be an important cellular mechanism that underlies the cognitive symptoms of depression.

animal behavior and cognition↗

Learning at variable attentional load requires cooperation between working memory, meta-learning and attention-augmented reinforcement learning

Flexible learning of changing reward contingencies can be realized with different strategies. A fast learning strategy involves using working memory of recently rewarded objects to guide choices. A slower learning strategy uses prediction errors to gradually update value expectations to improve choices. How the fast and slow strategies work together in scenarios with real-world stimulus complexity is not well known. Here, we disentangle their relative contributions in rhesus monkeys while they learned the relevance of object features at variable attentional load. We found that learning behavior across six subjects is consistently best predicted with a model combining (i) fast working memory (ii) slower reinforcement learning from differently weighted positive and negative prediction errors, as well as (iii) selective suppression of non-chosen feature values and (iv) a meta-learning mechanism adjusting exploration rates based on a memory trace of recent errors. These mechanisms cooperate differently at low and high attentional loads. While working memory was essential for efficient learning at lower attentional loads, enhanced weighting of negative prediction errors and meta-learning were essential for efficient learning at higher attentional loads. Together, these findings pinpoint a canonical set of learning mechanisms and demonstrate how they cooperate when subjects flexibly adjust to environments with variable real-world attentional demands. Significance statementLearning which visual features are relevant for achieving our goals is challenging in real-world scenarios with multiple distracting features and feature dimensions. It is known that in such scenarios learning benefits significantly from attentional prioritization. Here we show that beyond attention, flexible learning uses a working memory system, a separate learning gain for avoiding negative outcomes, and a meta-learning process that adaptively increases exploration rates whenever errors accumulate. These subcomponent processes of cognitive flexibility depend on distinct learning signals that operate at varying timescales, including the most recent reward outcome (for working memory), memories of recent outcomes (for adjusting exploration), and reward prediction errors (for attention augmented reinforcement learning). These results illustrate the specific mechanisms that cooperate during cognitive flexibility.

animal behavior and cognition↗

Fright and Fight: Role of predation and competition on mate search tactics of wild male zebrafish

Mate search tactics and association preferences among organisms in natural habitats can be dynamic and are determined by inherent trait preferences as well as the cost-benefit trade-offs associated with each mating decision. Two of the prime factors regulating mating decisions are the presence of competing conspecifics and predatory threats, both of which have important fitness consequences for the individual. We studied the influence of these two factors separately in mate search tactics and association preferences among zebrafish males. Male zebrafish were presented with a choice of two patches, consisting of different number of females, of which one patch was also associated with a predatory threat. We found that males made a preferential choice for the patch with more number of females only when the numerical difference between choices are starkly different, irrespective of the predatory threat associated with the patch. This points towards the role of numerical cognition in assessing cost-benefit tradeoffs in male zebrafish. We also studied the association preference of males in a multi-choice setup, consisting of four separate mixed-sex groups of zebrafish varying in densities. Our results showed that while test males preferred to visit the male-biased patches more often, they spent more time near female-biased patches or patches with equal sex ratio patches indicating the role of complex interplay of social cues in determining the associative behavior of males to a patch. This study, thus, sheds further light on the interactive roles of social cues and cognitive abilities in mate association patterns in this species.

animal behavior and cognition↗

Ornament, armament, or toolkit? Modelling how population size drives the evolution of birdsong, a functional cultural trait

Oscine songbirds have been an important study system for social learning, particularly because their learned songs provide an analog for human languages and music. Here we propose a different analogy; from an evolutionary perspective, could a birds song be more like an arrowhead than an aria? We modify a model of human tool evolution to accommodate cultural evolution of birdsong: each song learner chooses the most skilled available tutor to emulate, and each is more likely to produce an inferior copy than a superior one. Similarly to human tool evolution, we show that larger populations foster improvements in song over time, even when learners restrict their pool of tutors to a subset of individuals. We also demonstrate that songs could be simplified instead of lost after population bottlenecks if lower-quality traits are easier to imitate than higher-quality ones. We show that these processes could plausibly generate empirically observed patterns of song evolution, and we make predictions about the types of song elements most likely to be lost when populations shrink. More broadly, we aim to connect the modeling approaches used in human and non-human systems, moving toward a cohesive theoretical framework that accounts for both cognitive and demographic processes.

animal behavior and cognition↗

Undermatching is a consequence of policy compression

The matching law describes the tendency of agents to match the ratio of choices allocated to the ratio of rewards received when choosing among multiple options (Herrnstein, 1961). Perfect matching, however, is infrequently observed. Instead, agents tend to undermatch, or bias choices towards the poorer option. Overmatching, or the tendency to bias choices towards the richer option, is rarely observed. Despite the ubiquity of undermatching, it has received an inadequate normative justification. Here, we assume agents not only seek to maximize reward, but also seek to minimize cognitive cost, which we formalize as policy complexity (the mutual information between actions and states of the environment). Policy complexity measures the extent to which an agents policy is state-dependent. Our theory states that capacity-constrained agents (i.e., agents that must compress their policies to reduce complexity), can only undermatch or perfectly match, but not overmatch, consistent with the empirical evidence. Moreover, we validate a novel prediction about which task conditions exaggerate undermatching. Finally, we argue that a reduction in undermatching with higher dopamine levels in patients with Parkinsons disease is consistent with an increased policy complexity. Significance statementThe matching law describes the tendency of agents to match the ratio of choices allocated to different options to the ratio of reward received. For example, if option A yields twice as much reward as option B, matching states that agents will choose option A twice as much. However, agents typically undermatch: they choose the poorer option more frequently than expected. Here, we assume that agents seek to simultaneously maximize reward and minimize the complexity of their action policies. We show that this theory explains when and why undermatching occurs. Neurally, we show that policy complexity, and by extension undermatching, is controlled by tonic dopamine, consistent with other evidence that dopamine plays an important role in cognitive resource allocation.

animal behavior and cognition↗

Object permanence in domestic cats (Felis catus) using violation-of-expectancy by owner and stranger

Object permanence, the understanding that objects continue to exist while out of sight, has been extensively researched in human infants and animals. Cats have been shown to reach Stage 5 object permanence by passing successive visible displacement tests, but their understanding of Stage 6 object permanence is less clear. We tested 18 domestic cats on their understanding of Stage 4 (single visible displacement; SVD) and Stage 6 (invisible displacement; IVD) of object permanence in their home environment. Additionally, we investigated how person familiarity may influence study engagement. In SVD tests, the box manipulator (owner or researcher) hid a toy in one of two cardboard boxes for the cat to find. In IVD tests, we implemented a violation-of-expectancy procedure in which the box manipulator showed a toy re-appearing out of either a) the same box it was hidden in (consistent trials) or b) the box it was not hidden in (violation trials). Approximately half of the cats (56%) did not find the hidden toy in SVD trials, with 42% of these cats not attempting to find the toy, despite previous research demonstrating that cats can retrieve hidden objects in successive SVD tests. No predictors significantly influenced whether cats found the toy, or which box was checked first (toy or empty box). In IVD trials, we unexpectedly found that cats were more likely to play with and had longer gaze durations towards the toy and display more toy box-directed behaviours in consistent trials than violation trials. Similarly, we found that cats were more likely to display toy and empty box- directed behaviours in trials where the researcher acted as the box manipulator. Other factors such as breed, outdoor access, cat sex and the first person to act as the box manipulator also influenced toy-directed behaviours. We discuss the complexity of person familiarity in research contexts and highlight some methodological challenges in studying cat cognition.

animal behavior and cognition↗

Hornbills Adjust Response Speed According to Solvability of Patterned-Strings Problems

Cognitive adaptations for processing causal information are fundamental to flexible problem-solving across species. Patterned-string tasks offer a classic means-end problem, requiring subjects to identify the functional connection between an action and its distal outcome. This study investigated patterned-string pulling in East Asian hornbills, a cognitively underexplored avian taxon, to assess their problem-solving abilities and underlying cognitive mechanisms. Nine hornbills were presented with three string configurations of increasing complexity: parallel (baseline), broken (contact/no-contact), and crossed strings. Subjects reliably selected the correct string in the baseline and contact conditions, demonstrating sensitivity to physical continuity cues. In contrast, accuracy declined significantly in the crossed condition, which required recognition of spatial displacement between the baited and accessible string ends. Reaction time (RT) analyses revealed prolonged latencies in the crossed condition, likely reflecting increased cognitive demands or limited habituation. Individuals who failed often adopted suboptimal strategies, such as proximity or side biases. We also observed lateralisation effects, with a left-side RT advantage, suggesting hemispheric specialisation. These findings support the hypothesis that task complexity modulates both decision accuracy and response speed, highlight RT as a useful proxy for interim cognitive processing, and establish hornbills as a valuable comparative model for studying causal cognition in birds.

animal behavior and cognition↗