Journal of Experimental Psychology: General

Humans order numbers in space from left to right, with smaller quantities represented preferentially in the left hemispace and larger ones in the right hemispace. The direction of this mental number line (MNL), or more generally of number-space associations (NSA), is influenced by cultural habits such as reading and writing direction. However, a growing body of evidence from pre-verbal infants and non-human animals suggests that number-space mappings may also have biological foundations. In non-human primates, evidence for a directional MNL remains mixed, partly due to small sample sizes and methodological heterogeneity. Here, we tested samples of rhesus (Macaca mulatta) and crab-eating macaques (Macaca fascicularis) across two experiments using spontaneous food-related tasks. In Experiment 1, monkeys chose between identical food quantities (1x1 to 24x24) presented on the left and right. No systematic spatial choice bias emerged as a function of numerical magnitude, and hand use did not differ across exact numerical pairs, although exploratory analyses revealed magnitude-related modulations of manual responses. In Experiment 2, monkeys were habituated to small (4x4) or large (16x16) quantities and subsequently tested with the alternative quantity. Result showed significantly more leftward choices following numerical decreases (16[->]4) and more rightward choices following numerical increases (4[->]16), indicating that relative numerical context, rather than absolute magnitude, elicited directional spatial biases. These findings suggest that in macaques, number-space associations emerge most robustly in comparative contexts involving expectancy violations of magnitude.

Decisions based on evidence accumulated over time require rules governing when to end the accumulation process and commit to a choice. These rules control inherent trade-offs between decision speed and accuracy, which require careful balance to maximize quantities that depend on both like reward rate. We previously showed that, to maximize reward rate, normative decision rules adapt to changing task conditions (Barendregt et al., 2022). Here we used a novel task to examine whether and how people use adaptive rules for individual decisions under a variety of conditions, including changes in decision outcomes across trials and changes in evidence quality both across and within trials. We found that the participants tended to use rules that adjusted, at least partially, to predictable changes in task conditions to improve reward rate, consistent with a rationally bounded implementation of normative principles. These findings help inform our understanding of the extent and limits of flexible decision formation in the brain.

The ability to evaluate ones own knowledge states is often studied using paradigms in which participants make a decision and subsequently report their confidence. This structure has motivated hierarchical models in which confidence arises from a metacognitive process, distinct from the decision process itself, that estimates the probability that the choice is correct (Meyniel et al., 2015; Pouget et al., 2016; Fleming and Daw, 2017). Here, we contrast this framework with an alternative based on an intentional architecture (Shadlen et al., 2008). In this account, choice and confidence are determined simultaneously through a multidimensional drift-diffusion process, where each dimension represents one choice-confidence combination (Ratcliff and Starns, 2009, 2013). Choice, response time, and confidence jointly emerge when one of these accumulators reaches a decision bound. To adjudicate between these accounts, we fit both models to behavioral data from two perceptual tasks: a random-dots motion discrimination task with incentivized confidence reports, and a luminance discrimination task without feedback or incentives. The integrated model provided a superior fit for the incentivized motion task, whereas the hierarchical model more accurately captured behavior in the un-incentivized luminance task. These results suggest that confidence does not rely on a single computational mechanism, but rather its implementation may adapt to the specific demands and structure of the task.

Pretend play is a hallmark behavior in childhood where children create nonliteral meanings. Empirical data supporting the role of social cognition and the decoupling from literality are still scarce during early development. We explored here how the comprehension of pretense affects the visual exploratory behavior of toddlers (n = 44) and adults (n = 65) when they were exposed to short video clips in which an actress performed either real actions (e.g., eating jelly) or pretend actions (e.g., pretending to eat with imaginary food), while varying the complexity of those actions. We analyzed participants exploration of the face in the videos as exploitation of social information. We showed that all observers paid more attention to the face in pretend scenarios than in real ones, measured as longer total looking time in adults and more fixations and revisits to the face in both age groups. We also found more gaze shifts (a measure of information sampling) between the face and the moving hand in the pretend videos in both age groups, mainly at the initial stages of the actions. Additionally, analyses of the scanpaths structure using gaze entropy showed less order in the exploration of pretend videos in both age groups, suggesting that pretense involved greater uncertainty and increased information seeking. The less structured trajectories were observed again mainly in complex pretend scenarios. Taken together, our gaze results indicate that from its developmental origins, the comprehension of pretense relies on social processes linked with information seeking and exploration. Significance StatementDevelopmental theories have long debated whether pretend games are born in conjunction with social capacities in the second year or become integrated later in life. Our study shows that, much like adults, toddlers visually explore pretend scenes gathering more social information and in a less structured manner compared to real-world scenarios, suggesting that the emerging capacity to play with the meaning of things is linked with that of thinking of other minds early in life.

Spatialization in working memory refers to the spatial coding of non-spatial information along a mental horizontal line when encoding verbal material. This phenomenon is thought to support working memory by facilitating order encoding. Although it has been observed for both visually and auditorily presented stimuli, no direct comparison has yet examined whether these modalities rely on similar neural mechanisms. In this study, we investigated whether spatialization in visual and auditory modalities involves shared or distinct patterns of activity within the working-memory network. Forty-nine participants performed both a visual and an auditory working memory SPoARC task of the same verbal material, allowing to study the cortical patterns associated with distinct serial positions at both encoding and recognition across sensory modalities. Whole-brain analyses revealed similar frontoparietal networks across conditions. In addition, a representational similarity analysis (RSA) was conducted to assess the similarity of neural patterns between early and late serial positions in a sequence and across sensory modalities. This multivoxel pattern analysis revealed modality-dependent patterns distinguishing early and late positions in the inferior frontal gyrus. Additional modality-specific effects were observed in the anterior intraparietal sulcus in the visual modality and in the posterior hippocampus in the auditory modality. Drawing on the framework proposed by Bottini & Doeller (2020), we propose that order decoding in the IPS might reflect a low-dimensional spatial coding of order (e.g., along a horizontal axis), whereas order decoding in the hippocampus might reflect higher-dimensional spatial representations or temporal representations.

Despite growing evidence that many animals can evaluate quantities, the ecological relevance of numerical cognition remains debated, particularly outside vertebrates. Would individuals still rely on numerousness if less computationally demanding cues, visual features extracted at the early stage of visual processing, were available to assess quantity? In primates, individuals show a numerical bias as they tend to rely on the number of items rather than non-numerical cues, such as total area, to categorize quantities. In this study, we trained free-flying honeybees to discriminate between two and four items in conditions where numerosity covaried with the total area and perimeter (Experiment Size) or the convex hull (Experiment Space) cues, mimicking ecological contexts. Transfer tests assessed which numerical or non-numerical cues were learned and preferentially used by the bees. Bees primarily relied on numerousness over these non-numerical cues. Individual analyses revealed two consistent strategies: a "numerical bias" strategy, in which bees encoded numerical information while ignoring non-numerical cues, and a "generalist" strategy, where bees flexibly switched between cues and favored non-numerical information when cues conflicted. We further reported improved discrimination when smaller quantities appeared on the left and larger ones on the right, consistent with an oriented mental number line. Together, these findings demonstrate a spontaneous numerical bias in honeybees and reveal that individuals within the same species can adopt distinct strategies when evaluating quantity. Our findings also suggest that distantly related taxa like bees and primates may have independently evolved comparable mechanisms for quantity evaluation.

Balancing exploration and exploitation is a fundamental challenge for adaptive behavior, yet it remains unclear whether visual sampling and spatial locomotion reflect a single cross-domain trait or operate independently. We addressed this question by recording head-mounted eye-tracking and full-body motion tracking while 26 participants freely navigated "Westbrook", a large-scale virtual city for a total of 150 min across five sessions. From the movement trajectories we derived three spatial descriptors: median walking speed, occupancy entropy, and the proportion of explorative route choices. From the gaze data, we computed 38 robust visual descriptors encompassing fixation dynamics, pupil size, saccadic amplitude, gaze-head alignment, and transition entropy. Principal-component analysis reduced the visual descriptors to three components that captured 58 % of variance, with the first component (PC1) reflecting "gaze dynamism" (frequent shifts, larger saccades, higher transition entropy). Canonical correlation analysis revealed a strong coupling between spatial and visual behaviours: the first pair of canonical variates correlated at r = 0.68 (cross-validated r = 0.45), driven primarily by the association of high walking speed and occupancy entropy with elevated gaze dynamism. In contrast, the proportion of explorative route choices contributed little to this coupling. These findings demonstrate that individual differences in low-level locomotor speed and spatial coverage co-vary with an exploratory visual style, supporting the existence of a domain-general "exploration" factor that shapes both how people move through, and attend to, complex environments.

Practice enhances motor acuity, enabling movement execution with greater speed and accuracy. However, the learning principles underlying improvements in speed, accuracy, and efficiency remain less understood than those supporting motor skill acquisition and adaptation. Here, we examined motor execution in a skill-based practice task to characterize learning, retention, and generalization of motor acuity. Using a gamified two-dimensional racing task, right-handed participants controlled a stylus-driven car along a curved track as quickly and accurately as possible. Across two studies (N = 83 total, 54 females), participants completed 300 training laps on Session 1 and returned for Session 2 to assess retention and generalization to novel track configurations: one with altered spatial configuration (rotated track) and one requiring movement in the opposite direction of training (reverse track). Movement speed improved rapidly and showed robust, though incomplete, retention across sessions. Speed improvements generalized substantially to both novel tracks. Accuracy was high at training onset and showed strong retention. However, we do not observe offline gains between sessions. Notably, accuracy declined transiently for the novel track configurations, suggesting interference from prior training. Movement efficiency, indexed by path length, was retained and generalized to the rotated track. However, reversing movement direction impaired efficiency, revealing a movement direction effect. This effect persisted when training direction was reversed in a second study, with counterclockwise movements remaining slower and less efficient than clockwise movements. These findings show that practice produces durable and broadly transferable motor execution improvements, while inherent movement direction biases constrain how improvements generalize across contexts. New & NoteworthyThe learning principles underlying improvements in motor acuity remain less well understood than those governing other forms of motor learning. Prior work suggests that motor execution improvements show limited generalization. In contrast, the present findings demonstrate that execution-based practice can produce robust, transferable gains, while also revealing a key constraint: inherent movement direction biases that limit generalization. By characterizing learning, retention, and generalization, this work provides new insight into how motor acuity improvements compare with skill acquisition and adaptation.

A central question in psycholinguistics in visual word recognition is whether morphologically complex words are obligatorily decomposed into stems and affixes during visual word recognition or whether whole-word access can occur when forms are frequent and familiar. The present study investigated how morphological complexity and lexical frequency jointly shape neural responses by leveraging Korean nominal inflection, whose transparent stem-suffix structure permits a clean dissociation between base (stem) frequency and surface (whole-word) frequency. Twenty-five native Korean speakers completed a rapid event-related fMRI lexical decision task involving simple and inflected nouns that varied parametrically in both frequency measures. Representational similarity analysis (RSA) revealed robust encoding of surface frequency--but not base frequency--in the inferior frontal gyrus (IFG) pars opercularis and supramarginal gyrus (SMG), with significantly stronger correlations for inflected than simple nouns. Univariate analyses converged with this result: surface frequency selectively increased activation for inflected nouns in inferior parietal regions, whereas base frequency showed no reliable effects in any ROI. These findings challenge models positing obligatory pre-lexical decomposition, instead supporting accounts in which morphological processing is shaped by post-lexical, usage-driven lexical statistics. Taken together, our findings shed light on a distributed perspective on morphological processing, suggesting that structural and statistical factors jointly constrain access to morphologically complex forms.

Each individual person looks at natural scenes in their own unique way, resulting in a distinct perceptual experience of the world. However, little is known about why such differences in gaze emerge. Here, we test the hypothesis that idiosyncrasies in gaze behavior are predicted by inter-subject variations in internal models--expectations about how scenes typically look. In two experiments, we first characterized participants personal internal models by asking them to draw typical bathroom and kitchen scenes. Individual differences in these drawings were quantified using an objective deep learning pipeline and, in turn, related to individual differences in gaze behavior. In Experiment 1, where participants freely viewed a set of kitchen and bathroom photographs, inter-subject similarities in internal models did not predict inter-subject similarities in gaze. In Experiment 2, we encouraged strategic exploration through gaze-contingent viewing and a memory task. Here, inter-subject similarities in internal models predicted similarities in fixation frequency and the sequence in which different object categories were inspected. These findings suggest that the influence of internal models on visual exploration is stronger under increased sensory uncertainty and when expectation-guided sampling of the environment is encouraged. Together, our results provide new insights into how individual expectations shape gaze behavior and help explain why people differ in how they explore the visual world.

Brainstem arousal systems including the locus coeruleus noradrenaline system, re-spond transiently to behaviorally relevant events. Locus coeruleus activity also drives dilations of the pupil, which are often observed during cognitive tasks. The strength of pupil responses during encoding of stimulus material predicts the success of its later retrieval, which might reflect the impact of noradrenaline on synaptic plasticity and memory formation. The pupil also dilates in response to task-irrelevant sounds, which could therefore serve as a valuable tool for investigating causal effects of phasic, pupil-linked arousal on cognition. Here, we evaluated whether task-irrelevant white noise sounds affect memory formation and memory-based decisions. These sounds were played before, during or after the presentation of memoranda (images or spoken words). Memory success was measured in recognition and free recall tasks the day after. Trial-to-trial variations in the amplitude of pupil dilations during word encoding without task-irrelevant sounds predicted memory success. Task-irrelevant white-noise sounds also robustly dilated the pupil but did not improve memory formation for the words or the images. We conclude that pupil-linked arousal processes triggered by task-irrelevant sounds differ from those recruited endogenously during memory for-mation, for example in states of increased emotionality or attention.

Humanity has always admired and created artwork, but the neurocognitive mechanisms behind artistic experience are still elusive. Professional artists and their intimate relationship with their artworks provide a unique opportunity to study the nature of art experience due to their expertise in both art making and art appreciation. During two fMRI tasks, professional artists (N=20) made aesthetic judgments on their own and other artists paintings (aesthetic appreciation task); they also mentally reconstructed the moments when they conceived their artworks or, as a control condition, when they visited now-familiar places for the first time (reconstruction by imagery task). During art appreciation of their own (as compared to other artists) paintings, participants showed stronger recruitment of bilateral posterior parietal cortices, the left lateral occipitotemporal cortex, and the dorso-central sector of the right insula, that is, action-related brain regions also involved in encoding the emotional components of movements. The reconstruction of their own artistic creation (as compared to episodic memory retrieval) involved the left fronto-parietal network associated with motor cognition. Altogether, these results suggest that the mental representations of the actions involved in creating art are integral to the overall artistic experience of painters, supporting an embodied view of the artists experience of art.

Mental fatigue is known to impair cognitive and motor performance, but its impact on motor learning remains unclear. This study examined how mental fatigue affects skill acquisition in a sequential finger-tapping task. Twenty-eight participants were assigned to either a mental fatigue group, which completed a thirty-minute Stroop task, or a control group, which watched a documentary of equivalent duration. Both groups then trained on the finger-tapping task across multiple practice blocks with brief rest periods. Overall motor skill improved similarly in both groups. However, mental fatigue altered the pattern of acquisition: participants in the fatigue group showed decreased performance during practice blocks, which was compensated by larger gains during inter-block rest periods. A strong negative correlation was observed between online decrements and offline improvements, indicating that greater declines during practice were associated with larger gains during rest. This study highlights the critical role of rest periods in maintaining learning under cognitively demanding conditions and provides insight into how internal states, such as mental fatigue, can selectively influence the expression of performance without compromising overall learning.

How does the human brain represent the meaning of abstract symbols? Some theories postulate the existence of semantic spaces where concepts occupy positions that reflect their conceptual relationships. In the number domain, psychological evidence suggests that integers are represented along a mental number line which, with education, integrates higher-level number concepts such as fractions. To test this hypothesis, we recorded whole-brain 7T fMRI responses to integer and fraction symbols during a magnitude comparison task. Consistent with predictions, we found both behavioral and neural numerical distance effects. Activation vectors in intraparietal, inferior temporal, prefrontal, hippocampal, and parahippocampal cortices formed a two-dimensional semantic space organized by numerical magnitude and domain (fractions versus integers). Gaussian fits revealed a topographic map of numerical preferences in the anterior inferior parietal cortex, common to both domains. Our results suggest that, in educated adults, a joint semantic map integrates fractions and integers and supports symbolic magnitude representation and comparison.

Color is a prominent feature of visual experience, yet humans can recognize objects easily and accurately from grayscale images. We examined whether color becomes more useful when spatial information is degraded due to blurring. Participants viewed naturalistic scenes in color or grayscale, and reported whether a named target object was present across a range of blur levels that simulated optical defocus from 0-8 diopters. With unblurred images, performance did not differ between color and grayscale conditions, but as blur increased, recognition accuracy declined. Color provided a modest but reliable advantage at higher levels of blur, suggesting that color becomes increasingly useful when optical quality is degraded. We hypothesize that the evolutionary shift towards trichromacy may have been partially driven by the need to compensate for optical degradation due to aging and/or accumulated light exposure.

Some images are consistently remembered better than others, suggesting that memorability reflects intrinsic image properties. We tested whether within-category distinctiveness underlies this effect. Across three experiments (N = 477), participants categorized indoor scenes previously rated for subjective typicality and then completed recognition memory tests. Typical scenes were categorized faster and more accurately, but were remembered worse and showed a more liberal response bias than atypical scenes. These opposing effects were robust across categories. To link subjective typicality to visual representations, we quantified image distinctiveness using a convolutional neural network (CNN). Across layers, CNN-derived distinctiveness closely tracked human typicality judgments and predicted both categorization speed and memorability, with strongest effects in higher, semantic layers. Critically, the memory advantage for atypical scenes persisted even when most images were atypical, ruling out rarity within the experimental context. Together, the results show that intrinsic scene memorability reflects an images position within a category-specific representational space.

Play is a fundamental aspect of childhood and plays a crucial role in the development of creativity, yet its neural mechanisms remain poorly understood. We tested the hypothesis that more frequent play is associated with stronger functional integration among the default mode network (DMN), executive control network (CN), and salience network (SAL), as these cortical networks have been implicated in creativity in adults. In a preregistered study of infants and toddlers (Study 1; N = 143, 10 months-3 years, 67 boys, Baby Connectome Project), parent-reported play and imitation behaviors increased sharply from 1 to 2 years, and were associated with stronger within-DMN connectivity and DMN-CN coupling, controlling for age, sex, and head motion. In middle childhood (Study 2; N = 108, ages 4-11 years, 52 boys), parent-reported play frequency declined with age, as did cross-network coupling involving SAL. However, children who engaged more frequently in play showed higher DMN-SAL and CN-SAL connectivity. Finally, in a quasi-experimental comparison (Study 3; N = 45; ages 4-12 years, 20 boys), children enrolled in a curriculum that includes guided play (Montessori) showed higher DMN-SAL and DMN-CN connectivity than peers in traditional schools, suggesting that pedagogies that center child-led exploration might enable protracted brain network integration. Across these three studies, play was consistently associated with greater integration among DMN, SAL, and CN, a pattern previously linked to creativity in adults. Our findings offer a potential mechanism linking childhood play to later creativity through its role in supporting brain integration during development. Public Significant StatementO_LIPlay is widely believed to nurture childrens creativity, yet the brain mechanisms behind this link are not well understood. C_LIO_LIAcross three studies from infancy to middle childhood, we found that more frequent play was associated with stronger integration among brain networks tied to imagination, attention, and control. C_LIO_LIThese findings suggest that play may help build the neural foundation for later creative thinking. C_LI

The benefits of routines for daily functioning are widely acknowledged, yet, despite their apparent importance, methods for quantifying routine maintenance and the causes of their disruption remain lacking. Here, we propose a novel means of defining and quantifying routines (transition entropy). Using the transition entropy, we show that routines can be robustly elicited on tasks that require searching through a grid of squares for a hidden target. Over two experiments (N=100 each), we show that use of routines--as quantified by transition entropy--is robustly perturbed by frequent switches between search grids, as locations specific to the currently irrelevant grid become competitive for selection. Using a normative model that tracks task dynamics, we show that disruption to routines can be attributed to reduced sensitivity to the odds of success for completing a task. This suggests that routine maintenance may be disrupted by over-sensitivity to a lack of reward early in routine performance, or increased expectations regarding the utility of pursuing other tasks.

Binocular vision depends on the integration of matching visual features across the two eyes, while conflicting interocular signals can engage active inhibitory processes in the visual system. To investigate the temporal dynamics of these putative inhibitory processes, we examined how transitions between different binocular correlation states influence perceptual detectability and response speed. Using dynamic random-dot correlograms - free of monocular cues and allowing precise interocular manipulation - we presented brief target intervals embedded in longer background sequences. Stimuli varied in binocular correlation: correlated (C) patterns contained identical luminance profiles in both eyes, anticorrelated (A) patterns had inverted luminance dots, and uncorrelated (U) patterns had independent dot arrangements. Across three experiments, we measured (1) the presentation duration threshold required to detect a change in correlation, (2) simple reaction times (RTs) to the same transitions at suprathreshold levels, and (3) psychometric functions across durations for selected transitions. In Experiment 1, A[->]C transitions yielded significantly higher duration thresholds than C[->]A, indicating a suppressive influence associated with prior anticorrelation. In contrast, Experiment 2 showed that A[->]C transitions produced the shortest RTs, while C[->]U transitions were slowest, suggesting a rebound-like facilitation following prior suppression. Experiment 3 confirmed these temporal and contrast dependences, with opposite changes in contrast threshold and reaction times between transitions toward and away from the correlated fusional states. This divergence between perceptual onset and reaction time is consistent with a two-phase account in which binocular anticorrelation is associated with an initial suppressive phase followed by rebound-like facilitation that accelerates responses once the target becomes detectable. These findings are consistent with current models of binocular rivalry and fusion, and provide a temporally resolved behavioral perspective on how inhibitory control in sensory systems may dynamically influence subsequent responsiveness under conditions of perceptual ambiguity.

A critical aspect of human cognition is the ability to use our knowledge about the laws of physics to make predictions about physical events. Whether this ability is based on abstract processes or is grounded in our body-environment interactions remains an open debate. We used physical reasoning under altered gravity as a model system to show that humans real-time embodied experience modifies their high-level physical reasoning. Specifically, we tested participants in computerised reasoning games, while disrupting their gravitational signalling using Galvanic Vestibular Stimulation (GVS). Participants failed more and had suboptimal strategies under the GVS condition compared to no-GVS in games requiring reasoning about terrestrial gravity. However, the effects of GVS were reduced when the games included reasoning about altered gravity. Our findings demonstrate how the physical experience of the body shifts high-level cognitive skill as reasoning, suggesting that humans mental representation of the world is grounded in adaptable physical mechanisms.