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bioRxiv2025

Asian elephants (Elephas maximus) recognise human visual attention from body and face orientation

Visual attention has mostly been studied in primarily visual species, such as dogs and nonhuman primates. Although elephants rely more on acoustic and olfactory cues, they also use visual displays and gestures to communicate. Smet and Byrne (2014) showed that African savanna elephants (Loxodonta africana) recognise human visual attention based on face and body orientation, but this has not been investigated in Asian elephants (Elephas maximus). Here, we examined whether Asian elephants are sensitive to human attentional states. We tested ten captive female elephants in Thailand and analysed the frequency of experimenter-directed signals in a food-requesting task based on the experimenters body and face orientation. Elephants gestured significantly more when the experimenters body was oriented towards them than when it was turned away, and they gestured most frequently when her face was also oriented towards them. This suggests that body orientation may be the more salient visual cue, with face orientation contributing to the effect only when accompanied by body orientation. These findings show that Asian elephants, like African savanna elephants, understand the importance of visual attention for effective communication, contributing to our understanding of cognitive abilities across the elephant taxon and visual attention in animals.

animal behavior and cognition↗

bioRxiv2019

Long-term reduction of short wavelength light affects sustained attention and visuospatial working memory.

The short wavelength, i.e. blue light, is crucial for non-image forming effects such as entrainment of circadian system. Moreover, many studies showed that blue light enhances alertness and performance in cognitive tasks. However, most of scientific reports in this topic is based on studies using short exposure to blue or blue-enriched light and only few focused on the effects of its reduced transmittance, especially in longer period. The latter could potentially give insight into understanding if age-related sleep problems and cognitive decline are related to less amount of blue light reaching the retina, as our lenses become more yellow with age. In this study, we investigated the effects of prolonged blocking of blue light on cognitive functioning, in particular - sustained attention and visuospatial working memory, and sleep. We used amber contact lenses reducing transmittance of blue light by approximately 90% for the period of four weeks on a group of young, healthy participants. No changes were observed for measurements related to sleep and sleep-wake rhythm. The significant effect was shown both for sustained attention and visuospatial memory, i.e. the longer blocking the blue light lasted, the greater decrease in performance was observed. Additionally, the follow-up session was conducted (approximately one week after taking off the blue-blocking lenses) and revealed that in case of sustained attention this detrimental effect of blocking BL is fully reversible. Our findings provide evidence that prolonged reduction of BL exposure directly affects human cognitive functioning regardless of sleep-related conditions.

animal behavior and cognition↗

bioRxiv2019

Dissociating cholinergic influence on alertness andtemporal attention in primates in a simple reaction-timeparadigm

The ability to promptly respond to behaviourally relevant events depends on both general alertness and phasic changes in attentional state driven by temporal expectations. Using a variable foreperiod simple reaction time (RT) task in four adult male rhesus macaques, we investigated the role of the cholinergic system in alertness and temporal expectation. Foreperiod-effects on RT reflect temporal expectation, while alertness is quantified as overall response speed. We measured these RT parameters under vehicle treatment and systemic administration of the muscarinic receptor antagonist scopolamine. We also investigated whether and to what extent the effects of scopolamine were reversed by donepezil, a cholinesterase inhibitor widely used for the treatment of dementia. In the control condition, RT showed a continuous decrease as the foreperiod duration increased, which clearly indicated the effect of temporal expectation on RT. This foreperiod effect was mainly detectable on the faster tail of the RT distribution and was eliminated by scopolamine. Furthermore, scopolamine treatment slowed down the average RT. Donepezil treatment was efficient on the slower tail of the RT distribution and improved scopolamine-induced impairments only on the average RT reflecting a general beneficial effect on alertness without any improvement in temporal expectation. The present results highlight the role of the cholinergic system in temporal expectation and alertness in primates and help delineate the efficacy and scope of donepezil and other cholinomimetic agents as cognitive enhancers in present and future clinical practice.

animal behavior and cognition↗

bioRxiv2026

Multivariate Transfer Entropy Quantifies Information Transfer Between Dyadic Partners During Real-Time Perceptual Decision-Making

1Many social behaviours unfold continuously in time, yet most studies of social influence rely on discrete, trial-based paradigms. To quantify how humans actually integrate sensory and social signals during ongoing perceptual decision-making we here apply an information-theoretic analysis to data obtained with the Continuous Perceptual Report (CPR) paradigm. In the CPR task, pairs of participants tracked a dynamic random-dot motion stimulus, continuously reporting perceived direction and their confidence. Crucially, each participant had access to their partners moment-by-moment direction and confidence reports embedded within the stimulus display. We quantified information flow between the stimulus and behavioural responses using multivariate transfer entropy, allowing us to detect whether participants acquired additional information from their partner once stimulus information had been received and integrated it into their behavioural response. We show that real-time social interactions predominantly occur within matching behavioural dimensions: partners perceptual choices influence participants own choices, and partners confidence influences participants confidence. Using both, more stimulus and social information contributed to improved task performance, and participants selectively acquired more information from better-performing partners. Crucially, information integration was flexibly modulated by stimulus reliability: as sensory evidence became noisier, participants relied more on social information, but only when interacting with a highly reliable (e.g. computer-controlled) partner. Latency analyses further revealed that participants responded faster to changes in their partners perceptual choices of direction than to changes in stimulus direction. They also responded more slowly to changes in their partners confidence than to changes in their partners choice of direction. Together, these findings demonstrate that humans dynamically and adaptively integrate social information in a time-continuous manner, and they establish multivariate information-theoretic analyses as a powerful framework for studying real-time social cognition.

animal behavior and cognition↗

bioRxiv2023

A comparison of behavior paradigms assessing spatial memory in tree shrew

Spatial cognition enables animals to navigate the environment. Impairments in spatial navigation are often preclinical signs of Alzheimers disease (AD) in human. Therefore, evaluating spatial memory deficits is valuable when assessing incipient AD in animal models. The Chinese tree shrew, a close relative of primates, possesses many features that make it suitable for AD research. However, there is a scarcity of reliable behavior paradigms to monitor changes in spatial cognition in this species. To address this, we established reward-based paradigms in the radial-arm maze and the cheeseboard maze for tree shrew, and tested spatial memory of a group of twelve male animals in both tasks, along with a control water maze test, before and after bilateral lesions to the hippocampus, the brain region essential for spatial navigation. Tree shrews memorized target positions during training, and their task performance improved gradually until reaching a plateau in all three mazes. After the lesion, spatial learning was compromised in both newly-developed tasks, whereas memory retrieval was impaired in the water maze. Furthermore, individual task performance in both dry-land paradigms depended heavily on the size of remaining hippocampal tissue. Notably, all lesioned animals displayed spatial memory deficits in the cheeseboard task, but not in the other two paradigms. Our results suggest that the cheeseboard task currently represents the most sensitive paradigm for assessing spatial memory in tree shrew, with the potential to monitor progressive cognitive declines in aged or genetically modified animals developing AD-like symptoms. Significance StatementCognitive tests that monitor impairments in spatial memory play a crucial role in evaluating animal models with early-stage Alzheimers disease (AD). The Chinese tree shrew possesses many features suitable for an AD model, yet behavior tests assessing spatial cognition in this species are lacking. Here we developed novel behavior paradigms tailored to measure spatial memory in tree shrews and evaluated their sensitivity to changes in spatial learning by examining a group of hippocampus-lesioned animals. Our results indicate that the cheeseboard task effectively detects impairments in spatial memory and holds potential for monitoring the progressive cognitive decline in aged or genetically modified tree shrews that develop AD-like symptoms. This research may facilitate the use of tree shrew model in AD research.

animal behavior and cognition↗

bioRxiv2021

Evidence for a selective link between cooperation and individual recognition

The ability to recognize and discriminate among others is a frequent assumption of models of the evolution of cooperative behavior. At the same time, cooperative behavior has been proposed as a selective agent favoring the evolution of individual recognition abilities. While theory predicts that recognition and cooperation may co-evolve, data linking recognition abilities and cooperative behavior with fitness or evidence of selection are elusive. Here, we provide evidence of a fitness link between individual recognition and cooperation in the paper wasp Polistes fuscatus. Nest founding females in northern populations frequently form cooperative multiple foundress nests and possess highly variable facial patterns that mediate individual recognition. We describe a dearth of cooperative nesting, low phenotypic diversity, and a lack of individual recognition in southern populations. In a common garden experiment, northern co-foundress associations successfully reared offspring while all cooperative southern groups failed to rear any offspring, suggesting a fitness link between individual recognition and successful cooperation. Consistent with a selective link between individual recognition and cooperation, we find that rates of cooperative co-nesting correlate with identity-signaling color pattern diversity across the species range. Moreover, genomic evidence of recent positive selection on cognition loci likely to mediate individual recognition is substantially stronger in northern compared to southern P. fuscatus populations. Collectively, these data suggest that individual recognition and cooperative nesting behavior have co-evolved in P. fuscatus because recognition helps mediate conflict among co-nesting foundresses. This work provides evidence of a specific cognitive phenotype under selection because of social interactions, supporting the idea that social behavior can be a key driver of cognitive evolution.

animal behavior and cognition↗

bioRxiv2021

Constitutive depletion of brain serotonin differentially affects rats' social and cognitive abilities

BackgroundCentral serotonin is an essential neuromodulator for mental disorders. It appears a promising transdiagnostic marker of distinct psychiatric disorders and a common modulator of some of their key behavioral symptoms. We aimed to identify the behavioral markers of serotonergic function in rats and compare them to human deficits. MethodsWe applied a comprehensive profiling approach in adult male Tph2-/- rats constitutively lacking central serotonin. Under classical and ethological testing conditions, we tested each individuals cognitive, social and non-social abilities and characterized the group organization (i.e. social network, hierarchy). Using unsupervised machine learning, we identified the functions most dependent on central serotonin. ResultsIn classical procedures, Tph2-/- rats presented an unexpected normal cognitive profile. Under the complex and experimenter-free conditions of their home-cage, the same Tph2-/- rats presented drastic changes in their daily life. Brain serotonin depletion induced compulsive aggression and sexual behavior, hyperactive and hypervigilant stereotyped behavior, reduced self-care and body weight, and exacerbated corticosterone levels. Group-housed Tph2-/- rats showed strong social disorganization with disrupted social networks and hierarchical structure, which may arise from communication deficits and cognitive blunting. ConclusionsSerotonin depletion induced a profile reminiscent of the symptomatology of impulse control and anxiety disorders. Serotonin was necessary for behavioral adaptation to dynamic social environments. In classical testing conditions, our animal model challenged the concept of an essential role of serotonin in decision-making, flexibility, and impulsivity, although developmental compensations may have occurred. These contrasting findings highlight the need to generalize the evaluation of animal models multidimensional functions within the complexity of the social living environment.

animal behavior and cognition↗

bioRxiv2023

Differences Between Human and Non-Human Primate Theory of Mind: Evidence from Computational Modeling

Can non-human primates (NHPs) represent other minds? Answering this question is difficult because primates can fail tasks due to a lack of motivation or succeed through simpler strategies. Here we address these challenges through a computational theory-testing framework for NHP Theory of Mind. In this framework, each theory combines a proposed social representation with a parameter for how often it is used. This allow us to move beyond dichotomous positions about Theory of Minds presence or absence and instead analyze graded patterns of behavior as a combination of cognitive representations and their use. We apply this approach to one of the most foundational and well-studied aspects of Theory of Mind: the relation between seeing and knowing. Our results show that only theories in which NHPs have some representation of other minds can capture the qualitative pattern of successes and failures across five classic perspective-taking paradigms. However, these theories vary in their reliance on their representations, each showing significantly lower reliance than a human baseline. These results suggest that human and NHP social cognition differ in terms of reliance and possibly also in terms of representational complexity.

animal behavior and cognition↗

bioRxiv2024

The dentate gyrus provides flexibility for efficient spatial navigation

The hippocampus plays a critical role in spatial navigation and declarative memory. The dentate gyrus is the neurogenic region of the hippocampal formation and it has long been implicated in the fine separation of similar contexts or close object locations. However, it is unclear how an accurate discrimination could be beneficial to a goal-guided behavior in a changing environment. Therefore, we used chemogenetic inhibition to study the role of the dentate gyrus in a goal-guided spatial navigation paradigm over a familiar but dynamic crossword maze. Mice were challenged to localize a novel reward location from alternative pathways in two versions of the task with particular configurations in each experimental day. In the simple task, the two optimal paths to the goal shared some segments in their trajectory. In a more complex task, optimal trajectories demanded completely different directions to the reward location. Overall, mice with chemogenetic inhibition of the dentate gyrus were able to learn all the routes regardless the complexity of the task, similarly to control animals. However, after having solved a first route in the complex task, mice with dentate gyrus inhibition displayed an impairment to efficiently navigate over the alternate path. Our results demonstrate a role of the dentate gyrus in cognitive flexibility required to reach a goal in a changing familiar environment.

animal behavior and cognition↗

bioRxiv2021

No evidence for general intelligence in a fish

Differences in human general intelligence or reasoning ability can be quantified with the psychometric factor g, because individual performance across cognitive tasks is positively correlated. g also emerges in mammals and birds, is correlated with brain size and may similarly reflect general reasoning ability and behavioural flexibility in these species. To exclude the alternative that these positive cross-correlations may merely reflect the general biological quality of an organism or an inevitable by-product of having brains it is paramount to provide solid evidence for the absence of g in at least some species. Here, we show that wild-caught cleaner fish Labroides dimidiatus, a fish species otherwise known for its highly sophisticated social behaviour, completely lacks g when tested on ecologically non-relevant tasks. Moreover, performance in these experiments was not or negatively correlated with an ecologically relevant task, and in none of the tasks did fish caught from a high population density site outperform fish from a low-density site. g is thus unlikely a default result of how brains are designed, and not an automatic consequence of variation in social complexity. Rather, the results may reflect that g requires a minimal brain size, and thus explain the conundrum why the average mammal or bird has a roughly 10 times larger brain relative to body size than ectotherms. Ectotherm brains and cognition may therefore be organized in fundamentally different ways compared to endotherms.

animal behavior and cognition↗

bioRxiv2022

Olfactory detection and discrimination in domestic dogs (Canis lupus familiaris)

The extraordinary olfactory capabilities in detection and rescue dogs are well-known. However, the olfactory performance varies by breed and search environment (Jezierski et al., 2014), as well as by the quantity of training (Horowitz et al., 2013). While detection of an olfactory cue inherently demands a judgment regarding the presence or absence of a cue at a given location, olfactory discrimination requires an assessment of quantity, a task demanding more attention and, hence, decreasing reliability as an informational source (Horowitz et al., 2013). This study aims at gaining more clarity on detection and discrimination of olfactory cues in untrained dogs and in a variety of dog breeds. Using a two-alternative forced choice (2AFC) paradigm, we assessed olfactory detection scores by presenting a varied quantity of food reward under one or the other hidden cup, and discrimination scores by presenting two varied quantities of food reward under both hidden cups. We found relatively reliable detection performances across all breeds and limited discrimination abilities, modulated by breed. We discuss our findings in relation to the cognitive demands imposed by the tasks and the cephalic index of the dog breeds.

animal behavior and cognition↗

bioRxiv2022

General cognitive performance declines with female age and is negatively related to fledging success in a wild bird

Identifying the causes and fitness consequences of intraspecific variation in cognitive performance is fundamental to understand how cognition evolves. Selection may act on different cognitive traits separately or jointly as part of the general cognitive performance of the individual. To date, few studies have examined simultaneously whether individual cognitive performance covaries across different cognitive tasks, the relative importance of individual and social attributes in determining cognitive variation, and its fitness consequences in the wild. Here, we tested 38 wild southern pied babblers (Turdoides bicolor) on a cognitive test battery targeting associative learning, reversal learning and inhibitory control. We found that a single factor explained 59.5% of the variation in individual cognitive performance across tasks, suggestive of a general cognitive factor. General cognitive performance varied by age and sex; declining with age in females but not males. Older females also tended to produce a higher average number of fledglings per year compared to younger females. Analysing over 10 years of breeding data, we found that individuals with lower general cognitive performance produced more fledglings per year. Collectively, our findings support the existence of a trade-off between cognitive performance and reproductive success in a wild bird.

animal behavior and cognition↗

bioRxiv2023

Lasting dynamic effects of the psychedelic 2,5-dimethoxy-4-iodoamphetamine (+/-)-DOI) on cognitive flexibility

Psychedelic drugs can aid fast and lasting remission from various neuropsychiatric disorders, though the underlying mechanisms remain unclear. Preclinical studies suggest serotonergic psychedelics enhance neuronal plasticity, but whether neuroplastic changes can also be seen at cognitive and behavioural levels is unexplored. Here we show that a single dose of the psychedelic 2,5-dimethoxy-4-iodoamphetamine (({+/-})-DOI) affects structural brain plasticity and cognitive flexibility in young adult mice beyond the acute drug experience. Using ex vivo magnetic resonance imaging, we show increased volumes of several sensory and association areas one day after systemic administration of 2mgkg-1 ({+/-})-DOI. We then demonstrate lasting effects of ({+/-})-DOI on cognitive flexibility in a two-step probabilistic reversal learning task where 2mgkg-1 ({+/-})-DOI improved the rate of adaptation to a novel reversal in task structure occurring one-week post-treatment. Strikingly, ({+/-})-DOI-treated mice started learning from reward omissions, a unique strategy not typically seen in mice in this task, suggesting heightened sensitivity to previously overlooked cues. Crucially, further experiments revealed that ({+/-})-DOIs effects on cognitive flexibility were contingent on the timing between drug treatment and the novel reversal, as well as on the nature of the intervening experience. ({+/-})-DOIs facilitation of both cognitive adaptation and novel thinking strategies may contribute to the clinical benefits of psychedelic-assisted therapy, particularly in cases of perseverative behaviours and a resistance to change seen in depression, anxiety, or addiction. Furthermore, our findings highlight the crucial role of time-dependent neuroplasticity and the influence of experiential factors in shaping the therapeutic potential of psychedelic interventions for impaired cognitive flexibility.

animal behavior and cognition↗

bioRxiv2025

Behavioural flexibility and its drivers in semi-urban vervet monkeys

Phylogenetic comparisons suggest that behavioural flexibility facilitates success in urban environments. It remains less clear whether urbanization fosters cognitive skills that require flexibility, or whether successful individuals in urban environments simply apply pre-evolved skills to solve new problems. To investigate whether variation in anthropogenic experience drives behavioural flexibility required to solve a new technical problem, we presented 42 semi-urban vervet monkeys (Chlorocebus pygerythrus) with a three-phase foraging experiment. For this, we measured each monkeys tendency to "raid" human structures in their search for food to assess whether their anthropogenic experience explained individual variation in flexibility and problem-solving skills. We then used a probabilistic model to describe the series of monkeys actions when attempting to solve the foraging experiment. This allowed us to quantify three traits encompassing individuals behavioural flexibility: switch tendency between solutions, innovativeness, and learning sensitivity. We found that individuals switch tendency and innovativeness were not explained by anthropogenic experience and that the three traits were unrelated to each other. Moreover, neither switch tendency, nor innovativeness predicted the monkeys human food consumption in this habitat. Thus, our study suggests that behavioural flexibility is not driven by anthropogenic foraging experience. Contrasting with the hypothesis that urbanization selects for behavioural flexibility, our findings instead imply that vervet monkeys already had the sufficient behavioural flexibility and cognitive capacities to successfully exploit the urban habitat.

animal behavior and cognition↗

bioRxiv2025

Casual Versus Fractal Modularity in Mammalian Social Organisation

Multilevel social systems in birds and mammals have attracted considerable interest, not least because they share some similarities with the natural structural organisation of human societies. I use a fractal analysis of a taxonomically wide range of species reported to have multilevel social systems to show that these in fact involve two very different phenomena: those that are fractally modular and those that are casually modular. The first have a strictly fractal structure identical to that found in human societies; the second lack formal structural coherence and are closer in form to casual herds. The former are associated with larger neocortices than the latter, suggesting that they may be cognitively more demanding. Understanding the evolution of multilevel systems and the cognition that underpins them obliges us to adopt a more nuanced approach to social evolution.

animal behavior and cognition↗

bioRxiv2025

Learning and Memory Performance in a Rat Model of Perigestational Opioid Exposure

Rationale: Nearly one third of women of reproductive age in the United States are prescribed opioids annually; 14% will fill an opioid prescription during pregnancy, and one in five report misuse. Opioid use during pregnancy has given rise to an increasing population of infants born with gestational opioid exposure. At school age, these children show neurodevelopmental impairment and higher rates of learning disability. Objectives: To characterize how exposure to exogenous opioids during brain development affects learning and memory performance, our lab has developed a rat model of perigestational opioid exposure that closely recapitulates a clinically relevant dosing timeline by beginning morphine exposure before pregnancy and continuing through the first postnatal week. Male and female offspring generated from this model were assessed on several facets of learning and memory during adolescence. Results: Here, we report that morphine exposure selectively impaired spatial learning, as measured by the Barnes Maze, and associative learning, as measured by an Attentional Set Shift Task, without significantly impacting working or short-term memory. Environmental enrichment rescued the spatial learning deficit in males but not females. Conclusion: Our findings suggest that gestational opioid exposure can impair performance in complex cognitive tasks that are attenuated by non-invasive, non-pharmacological intervention.

animal behavior and cognition↗

bioRxiv2026

Japanese quail (Coturnix japonica) selected for high social motivation rely on conspecifics for buffering but not learning in sociocognitive tasks

Social living is widely considered a key driver of cognitive evolution, yet individuals within a species and even within the same group can differ substantially in their sociability (i.e., an individuals propensity to form and maintain social bonds), which can ultimately shape the social environment they experience by influencing how they interact with, respond to, and engage in it. How such individual variation in this personality trait affects social cognition, however, remains poorly understood. To address this question, we used two selectively bred lines of Japanese quail (Coturnix japonica) that consistently differ in social motivation, a key component of sociability, which we used as a proxy for this broader trait. In these lines, S+ individuals show high social motivation, whereas S- individuals show low social motivation. We compared their sociocognitive performance across three tasks: a gaze following task, a social buffering task, and a social discrimination learning task. Our findings revealed that Japanese quail reliably followed the gaze of conspecifics, providing the first evidence of this ability in this species. However, there was no difference between lines, suggesting that basic gaze following into the distance is independent of social motivation. In contrast, line differences emerged in the other tasks. S+ quail were more sensitive and less adaptable in response to environmental changes, particularly under social isolation, although the presence of a conspecific strongly buffered these effects. S- quail, on the other hand, outperformed S+ individuals in the social discrimination learning task, rapidly exploiting available social cues to guide foraging decisions. Overall, this study demonstrates that social motivation modulates sociocognitive performance in a context-dependent manner. Rather than conferring a general cognitive advantage, divergent selection on social motivation leads to different strategies of social information use across tasks and contexts, highlighting the complex interplay between personality, social environment, and cognition.

animal behavior and cognition↗

bioRxiv2026

Investigating the Effects of Psilocybin on Cognitive Flexibility in Touchscreen and Naturalistic Variations of the Probabilistic Reversal Learning Task

RationalePsychedelic compounds such as psilocybin have attracted growing interest for their potential therapeutic effects in psychiatric disorders, with improvements in cognitive flexibility proposed as a possible mechanism of action. However, the effects of psychedelics on cognitive flexibility remain poorly understood. ObjectiveThis study aimed to examine the acute and post-acute effects of psilocybin (0.1, 0.3, 1 mg/kg) and lysergic acid diethylamide (LSD, (0.02, 0.04, 0.08 mg/kg) on cognitive flexibility in male rats. MethodsThis was tested using two variants of the probabilistic reversal learning task (PRLT): a touchscreen-based operant task and a more ethological foraging-based task. ResultsIn the touchscreen PRLT, acute psilocybin disrupted task engagement, with animals completing fewer trials and showing increased trial initiation latency, although psilocybin also showed a trend toward faster initial rule acquisition. However, psilocybin did not significantly alter the number of rule changes achieved, a canonical measure of cognitive flexibility, or feedback sensitivity. LSD similarly produced limited acute effects, although the highest dose reduced lose-shift probability, suggesting decreased sensitivity to negative feedback under some conditions. Post-acute effects of psilocybin were minimal in both PRLT variants and, where LSD effects were observed these occurred across different doses and timepoints without a consistent pattern. ConclusionsOverall, these findings suggest that serotonergic psychedelics do not robustly enhance reversal learning in these paradigms and that apparent learning effects may reflect transient disruptions in task engagement rather than improvements in cognitive flexibility. These results also highlight potential limitations of these PRLT paradigms for detecting psychedelic-induced changes in cognitive flexibility in rodents.

animal behavior and cognition↗