Weaker number sense accounts for impaired numerosity perception in dyscalculia: Behavioral and computational evidence.

Impaired numerosity perception in developmental dyscalculia (low "number acuity") has been interpreted as evidence of reduced representational precision in the neurocognitive system supporting non-symbolic number sense. However, recent studies suggest that poor numerosity judgments might stem from stronger interference from non-numerical visual information, in line with alternative accounts that highlight impairments in executive functions and visuospatial abilities in the etiology of dyscalculia. To resolve this debate, we used a psychophysical method designed to disentangle the contribution of numerical and non-numerical features to explicit numerosity judgments in a dot comparison task and we assessed the relative saliency of numerosity in a spontaneous categorization task. Children with dyscalculia were compared to control children with average mathematical skills matched for age, IQ, and visuospatial memory. In the comparison task, the lower accuracy of dyscalculics compared to controls was linked to weaker encoding of numerosity, but not to the strength of non-numerical biases. Similarly, in the spontaneous categorization task, children with dyscalculia showed a weaker number-based categorization compared to the control group, with no evidence of a stronger influence of non-numerical information on category choice. Simulations with a neurocomputational model of numerosity perception showed that the reduction of representational resources affected the progressive refinement of number acuity, with little effect on non-numerical bias in numerosity judgments. Together, these results suggest that impaired numerosity perception in dyscalculia cannot be explained by increased interference from non-numerical visual cues, thereby supporting the hypothesis of a core number sense deficit. RESEARCH HIGHLIGHTS: A strongly debated issue is whether impaired numerosity perception in dyscalculia stems from a deficit in number sense or from poor executive and visuospatial functions. Dyscalculic children show reduced precision in visual numerosity judgments and weaker number-based spontaneous categorization, but no increasing reliance on continuous visual properties. Simulations with deep neural networks demonstrate that reduced neural/computational resources affect the developmental trajectory of number acuity and account for impaired numerosity judgments. Our findings show that weaker number acuity in developmental dyscalculia is not necessarily related to increased interference from non-numerical visual cues.

The interplay between spatial and non-spatial grouping cues over approximate number perception.

Humans and animals share the cognitive ability to quickly extract approximate number information from sets. Main psychophysical models suggest that visual approximate numerosity relies on segmented units, which can be affected by Gestalt rules. Indeed, arrays containing spatial grouping cues, such as connectedness, closure, and even symmetry, are underestimated compared to ungrouped arrays with equal low-level features. Recent evidence suggests that non-spatial cues, such as color-similarity, also trigger numerosity underestimation. However, in natural vision, several grouping cues may coexist in the scene. Notably, conjunction of grouping cues (color and closure) reduces perceived numerosity following an additive rule. To test whether the conjunction-effect holds for other Gestalt cues, we investigated the effect of connectedness and symmetry over numerosity perception both in isolation and, critically, in conjunction with luminance similarity. Participants performed a comparison-task between a reference and a test stimulus varying in numerosity. In Experiment 1, test stimuli contained two isolated groupings (connectedness or luminance), a conjunction (connectedness and luminance), and a neutral condition (no groupings). Results show that point of subjective equality was higher in both isolated grouping conditions compared to the neutral condition. Furthermore, in the conjunction condition, the biases from isolated grouping cues added linearly, resulting in a numerosity underestimation equal to the sum of the isolated biases. In Experiment 2 we found that conjunction of symmetry and luminance followed the same additive rule. These findings strongly suggest that both spatial and non-spatial isolated cues affect numerosity perception. Crucially, we show that their conjunction effect extends to symmetry and connectedness.

Developmental changes of the impact of visual cues on ANS acuity across grades 1-5: Different patterns of visual cues on numerosity processing.

Numerous studies have consistently demonstrated the presence of the approximate number system (ANS) throughout development. Research has also revealed that visual cues may influence the ANS acuity, which may change with age. However, most studies have drawn conclusions based on performance differences between incongruent and congruent trials, which may be confounded by an individual's ability to inhibit interference. Therefore, to examine the developmental changes of the impact of visual cues on ANS acuity, we utilized congruent trials with varying visual cues. Our sample comprised Chinese children from grade one to grade five. We manipulated the salience of numerical cues (numerical ratio) and visual cues (dot size) in a non-symbolic numerosity comparison task. The results revealed a discernible leap in development from first to third grade and first to fifth grade; however, this upward trajectory did not persist into the transition from third to fifth grade, where no appreciable advancement was observed. Moreover, we observed different effects of visual cues on the dot-comparison task depending on the numerical cues and age. Specifically, visual cues (i.e., dot size) only facilitated ANS acuity in older school-aged children when numerical cues were weakened. The results indicate the presence of two distinct magnitude representational systems-one for the numerical dimension and another for the non-numerical dimension-during development.

Numbers in Context: Cardinals, Ordinals, and Nominals in American English.

There are three main types of number used in modern, industrialized societies. Cardinals count sets (e.g., people, objects) and quantify elements of conventional scales (e.g., money, distance), ordinals index positions in ordered sequences (e.g., years, pages), and nominals serve as unique identifiers (e.g., telephone numbers, player numbers). Many studies that have cited number frequencies in support of claims about numerical cognition and mathematical cognition hinge on the assumption that most numbers analyzed are cardinal. This paper is the first to investigate the relative frequencies of different number types, presenting a corpus analysis of morphologically unmarked numbers (not, e.g., "eighth" or "21st") in which we manually annotated 3,600 concordances in the Corpus of Contemporary American English. Overall, cardinals are dominant-both pure cardinals (sets) and measurements (scales)-except in the range 1,000-10,000, which is dominated by ordinal years, like 1996 and 2004. Ordinals occur less often overall, and nominals even less so. Only for cardinals do round numbers, associated with approximation, dominate overall and increase with magnitude. In comparison with other registers, academic writing contains a lower proportion of measurements as well as a higher proportion of ordinals and, to some extent, nominals. In writing, pure cardinals and measurements are usually represented as number words, but measurements-especially larger, unround ones-are more likely to be numerals. Ordinals and nominals are mostly represented as numerals. Altogether, this paper reveals how numbers are used in American English, establishing an initial baseline for any analyses of number frequencies and shedding new light on the cognitive and psychological study of number.

Rational number representation by the approximate number system.

The approximate number system (ANS) enables organisms to represent the approximate number of items in an observed collection, quickly and independently of natural language. Recently, it has been proposed that the ANS goes beyond representing natural numbers by extracting and representing rational numbers (Clarke & Beck, 2021a). Prior work demonstrates that adults and children discriminate ratios in an approximate and ratio-dependent manner, consistent with the hallmarks of the ANS. Here, we use a well-known "connectedness illusion" to provide evidence that these ratio-dependent ratio discriminations are (a) based on the perceived number of items in seen displays (and not just non-numerical confounds), (b) are not dependent on verbal working memory, or explicit counting routines, and (c) involve representations with a part-whole (or subset-superset) format, like a fraction, rather than a part-part format, like a ratio. These results vindicate key predictions of the hypothesis that the ANS represents rational numbers.

Sensorimotor mechanisms selective to numerosity derived from individual differences.

We have previously shown that after few seconds of adaptation by finger-tapping, the perceived numerosity of spatial arrays and temporal sequences of visual objects displayed near the tapping region is increased or decreased, implying the existence of a sensorimotor numerosity system (Anobile et al., 2016). To date, this mechanism has been evidenced only by adaptation. Here, we extend our finding by leveraging on a well-established covariance technique, used to unveil and characterize 'channels' for basic visual features such as colour, motion, contrast, and spatial frequency. Participants were required to press rapidly a key a specific number of times, without counting. We then correlated the precision of reproduction for various target number presses between participants. The results showed high positive correlations for nearby target numbers, scaling down with numerical distance, implying tuning selectivity. Factor analysis identified two factors, one for low and the other for higher numbers. Principal component analysis revealed two bell-shaped covariance channels, peaking at different numerical values. Two control experiments ruled out the role of non-numerical strategies based on tapping frequency and response duration. These results reinforce our previous reports based on adaptation, and further suggest the existence of at least two sensorimotor number channels responsible for translating symbolic numbers into action sequences.

Understanding the social-emotional components of our "number sense": insights from a novel non-symbolic numerical comparison task.

Introduction: A large body of work has identified a core sense of number supported by the Approximate Number System (ANS) that is present in infancy and across species. Although it is commonly assumed that the ANS directly processes perceptual input and is relatively independent from affective factors, some evidence points at a correlation between ANS performance and math anxiety. However, the evidence is mixed. We tested whether giving participants active control in completing a numerical task would change the relationship between math anxiety on performance. Methods: Adult participants (N = 103) completed a novel four-alternative-forced-choice non-symbolic numerical comparison task. In a repeated-measures design, participants either passively viewed different dot arrays or actively chose to view each array (i.e., active information-seeking) before deciding on the largest quantity. Participants also provided confidence judgments during the passive version of the task. Results: We replicated the ratio-dependent signature in participants' accuracy in both the passive and active versions of the task using this novel paradigm, as well as in trial-level confidence judgments and information-seeking behavior. Participants' self-reported math anxiety significantly correlated with their accuracy on the passive version of the task. Critically, the correlation disappeared in the active version of the task. Gender also emerged as a predictor of confidence judgments and a moderator of the effect of task on overall accuracy and the effect of active information seeking on accuracy in the active version of the task. Exploratory analysis of estimated Weber Fraction suggests that these results may be driven by auxiliary factors instead of changes in ANS acuity. Conclusion: These findings have implications for understanding the relationship between math anxiety and performance on numerical tasks.

On the functional independence of numerical acuity and visual working memory.

Deciding where to direct our vehicle in a crowded parking area or where to line up at an airport gateway relies on our ability to appraise the numerosity of multitudes at a glimpse and react accordingly. Approximating numerosities without actually counting is an ontogenetically and phylogenetically primordial ability, given its presence in human infants shortly after birth, and in primate and non-primate animal species. Prior research in the field suggested that numerosity approximation is a ballistic automatism that has little to do with human cognition as commonly intended. Here, we measured visual working memory capacity using a state-of-the-art change detection task and numerosity approximation using a dot-comparison task, and found a null correlation between these two parametrical domains. By checking the evidential strength of the tested correlation using both classic and Bayesian analytical approaches, as well as the construct validity for working memory capacity and numerosity approximation estimates, we concluded that the present psychophysical evidence was sufficiently strong to support the view that visual working memory and numerosity approximation are likely to rely on functionally independent stages of processing of the human cognitive architecture.

Modeling Magnitude Discrimination: Effects of Internal Precision and Attentional Weighting of Feature Dimensions.

Given a rich environment, how do we decide on what information to use? A view of a single entity (e.g., a group of birds) affords many distinct interpretations, including their number, average size, and spatial extent. An enduring challenge for cognition, therefore, is to focus resources on the most relevant evidence for any particular decision. In the present study, subjects completed three tasks-number discrimination, surface area discrimination, and convex hull discrimination-with the same stimulus set, where these three features were orthogonalized. Therefore, only the relevant feature provided consistent evidence for decisions in each task. This allowed us to determine how well humans discriminate each feature dimension and what evidence they relied on to do so. We introduce a novel computational approach that fits both feature precision and feature use. We found that the most relevant feature for each decision is extracted and relied on, with minor contributions from competing features. These results suggest that multiple feature dimensions are separately represented for each attended ensemble of many items and that cognition is efficient at selecting the appropriate evidence for a decision.

Humans can sense large numbers of objects in a box by touch alone.

Everyday experiences suggest that a container, such as a box of cereal, can convey pertinent information about the nature and quantity of its content. This study investigated how well people can judge large quantities of objects in a container through haptic perception. Stimuli consisted of plastic drinking straws cut to "small" (1.5 cm) or "big" (4.5 cm) pieces contained in plastic food containers. Participants performed both a magnitude estimation of the number of objects and a direct estimation of the proportion of the container perceived to be filled with objects. Overall, participants demonstrated considerable accuracy for both tasks and irrespective of the size of the content. Post-experiment interviews revealed three potential strategies. Participants either focused on the container's contents, the excess space in the container, or the perceived weight of the container (content).

Corrigendum: Effects of intelligence and approximate number system on the non-symbolic division ability in preschoolers.

[This corrects the article DOI: 10.3389/fpsyg.2023.961140.].

The Interaction between Congruency and Numerical Ratio Effects in the Nonsymbolic Comparison Test.

The nonsymbolic comparison task is used to investigate the precision of the Approximate Number Sense, the ability to process discrete numerosity without counting and symbols. There is an ongoing debate regarding the extent to which the ANS is influenced by the processing of non-numerical visual cues. To address this question, we assessed the congruency effect in a nonsymbolic comparison task, examining its variability across different stimulus presentation formats and numerical proportions. Additionally, we examined the variability of the numerical ratio effect with the format and congruency. Utilizing generalized linear mixed-effects models with a sample of 290 students (89% female, mean age 19.33 years), we estimated the congruency effect and numerical ratio effect for separated and intermixed formats of stimulus presentation, and for small and large numerical proportions. The findings indicated that the congruency effect increased in large numerical proportion conditions, but this pattern was observed only in the separated format. In the intermixed format, the congruency effect was insignificant for both types of numerical proportion. Notably, the numerical ratio effect varied for congruent and incongruent trials in different formats. The results may suggest that the processing of visual non-numerical parameters may be crucial when numerosity processing becomes noisier, specifically when numerical proportion becomes larger. The implications of these findings for refining the ANS theory are discussed.

Development of precision of non-symbolic arithmetic operations in 4-6-year-old children.

Children can represent the approximate quantity of sets of items using the Approximate Number System (ANS), and can perform arithmetic-like operations over ANS representations. Previous work has shown that the representational precision of the ANS develops substantially during childhood. However, less is known about the development of the operational precision of the ANS. We examined developmental change in the precision of the solutions to two non-symbolic arithmetic operations in 4-6-year-old U.S. children. We asked children to represent the quantity of an occluded set (Baseline condition), to compute the sum of two sequentially occluded arrays (Addition condition), or to infer the quantity of an addend after observing an initial array and then the array incremented by the unknown addend (Unknown-addend condition). We measured the precision of the solutions of these operations by asking children to compare their solutions to visible arrays, manipulating the ratio between the true quantity of the solution and the comparison array. We found that the precision of ANS representations that were not the result of operations (in the Baseline condition) was higher than the precision of solutions to ANS operations (in the Addition and Unknown-addend conditions). Further, we found that precision in the Baseline and Addition conditions improved significantly between 4 and 6 years, while precision in the Unknown-Addend condition did not. Our results suggest that ANS operations may inject "noise" into the representations they operate over, and that the development of the precision of different operations may follow different trajectories in childhood.

The role of non-numerical information in the perception of temporal numerosity.

Numerosity perception refers to the ability to make rapid but approximate estimates of the quantity of elements in a set (spatial numerosity) or presented sequentially (temporal numerosity). Whether numerosity is directly perceived or indirectly recomputed from non-numerical features is a highly debated issue. In the spatial domain, area and density have been suggested as the main parameters through which numerosity would be recomputed. In the temporal domain, stimuli duration and temporal frequency could be similarly exploited to retrieve numerosity. By adapting a psychophysical technique previously exploited in the spatial domain, we investigated whether temporal visual numerosity is directly perceived. Adult participants observed sequences of visual impulses sampled from a stimulus space spanning several levels of temporal frequency and duration (and hence numerosity), and then reproduced the sequence as accurately as possible via a series of keypresses. Crucially, participants were not asked to reproduce any particular property (such as number of impulses) but were free to choose any available cue (such as total duration, or temporal frequency). The results indicate that while the overall sequence duration was barely considered, numerosity and temporal frequency were both spontaneously used as the main cues to reproduce the sequences, with a slight but significant dominance of numerosity. Overall, the results are in line with previous literature suggesting that numerosity is directly encoded, even for temporal sequences, but a non-numerical feature (temporal frequency) is also used in reproducing sequences.

Effects of intelligence and approximate number system on the non-symbolic division ability in preschoolers.

Recently, it has become evident that cognitive abilities such as the approximate number system (ANS), number knowledge, and intelligence affect individuals' fundamental mathematical ability. However, it is unclear which of these cognitive abilities have the greatest impact on the non-symbolic division ability in preschoolers. Therefore, in the present study, we included 4- to 6-year-old Korean preschoolers without prior formal education of division in order to test their ability to solve non-symbolic division problems, ANS acuity, and intelligence, and to determine the interrelationships among those functions (N = 38). We used the Panamath Dot Comparison Paradigm to measure the ANS acuity, employed non-symbolic division tasks to measure the ability to solve non-symbolic division problems, and measured the intelligence using the Korean version of the WPPSI-IV (Wechsler Preschool Primary Scale of Intelligence-IV). Our results showed that, in all conditions of the non-symbolic division tasks, the 4- to 6-years old children were able to perform better than chance level. Additionally, in a relatively easy condition, the children's performance showed a significant positive correlation with full-scale intelligence quotient (FSIQ) and ANS acuity; however, in a more complex condition, only FSIQ was significantly correlated with their performance. Overall, we found significant relationships between the children's performance in the non-symbolic division tasks and verbal comprehension, fluid reasoning, and processing speed index. Taken together, our findings demonstrate that preschoolers without formal education on the arithmetic problem solving can solve non-symbolic division problems. Moreover, we suggest that both FSIQ and ANS ability play essential roles in children's ability to solve non-symbolic division problems, highlighting the significance of intelligence on children's fundamental mathematical ability.

Fundamental units of numerosity estimation.

Humans can approximately enumerate a large number of objects at a single glance. While several mechanisms have been proposed to account for this ability, the fundamental units over which they operate remain unclear. Previous studies have argued that estimation mechanisms act only on topologically distinct units or on units formed by spatial grouping cues such as proximity and connectivity, but not on units grouped by similarity. Over four experiments, we tested this claim by systematically assessing and demonstrating that similarity grouping leads to underestimation, just as spatial grouping does. Ungrouped objects with the same low-level properties as grouped objects did not cause underestimation. Further, the underestimation caused by spatial and similarity grouping was additive, suggesting that these grouping processes operate independently. These findings argue against the proposal that estimation mechanisms operate solely on topological units. Instead, we conclude that estimation processes act on representations constructed after Gestalt grouping principles, whether similarity based or spatial, have organised incoming visual input.

Numerical discrimination in Drosophila melanogaster.

Sensitivity to numbers is a crucial cognitive ability. The lack of experimental models amenable to systematic genetic and neural manipulation has precluded discovering neural circuits required for numerical cognition. Here, we demonstrate that Drosophila flies spontaneously prefer sets containing larger numbers of objects. This preference is determined by the ratio between the two numerical quantities tested, a characteristic signature of numerical cognition across species. Individual flies maintained their numerical choice over consecutive days. Using a numerical visual conditioning paradigm, we found that flies are capable of associating sucrose with numerical quantities and can be trained to reverse their spontaneous preference for large quantities. Finally, we show that silencing lobula columnar neurons (LC11) reduces the preference for more objects, thus identifying a neuronal substrate for numerical cognition in invertebrates. This discovery paves the way for the systematic analysis of the behavioral and neural mechanisms underlying the evolutionary conserved sensitivity to numerosity.

Is Nonsymbolic Arithmetic Truly "Arithmetic"? Examining the Computational Capacity of the Approximate Number System in Young Children.

Young children with limited knowledge of formal mathematics can intuitively perform basic arithmetic-like operations over nonsymbolic, approximate representations of quantity. However, the algorithmic rules that guide such nonsymbolic operations are not entirely clear. We asked whether nonsymbolic arithmetic operations have a function-like structure, like symbolic arithmetic. Children (n = 74 4- to -8-year-olds in Experiment 1; n = 52 7- to 8-year-olds in Experiment 2) first solved two nonsymbolic arithmetic problems. We then showed children two unequal sets of objects, and asked children which of the two derived solutions should be added to the smaller of the two sets to make them "about the same." We hypothesized that, if nonsymbolic arithmetic follows similar function rules to symbolic arithmetic, then children should be able to use the solutions of nonsymbolic computations as inputs into another nonsymbolic problem. Contrary to this hypothesis, we found that children were unable to reliably do so, suggesting that these solutions may not operate as independent representations that can be used inputs into other nonsymbolic computations. These results suggest that nonsymbolic and symbolic arithmetic computations are algorithmically distinct, which may limit the extent to which children can leverage nonsymbolic arithmetic intuitions to acquire formal mathematics knowledge.

Electrophysiological Comparison of Cumulative Area and Non-Symbolic Number Judgments.

Despite the importance of representing different magnitudes (i.e., number and cumulative area) for action planning and formal mathematics, there is much debate about the nature of these representations, particularly the extent to which magnitudes interact in the mind and brain. Early interaction views suggest that there are shared perceptual processes that form overlapping magnitude representations. However, late interaction views hold that representations of different magnitudes remain distinct, interacting only when preparing a motor response. The present study sheds light on this debate by examining the temporal onset of ratio and congruity effects as participants made ordinal judgments about number and cumulative area. Event-related potentials (ERPs) were recorded to identify whether the onset of such effects aligned with early versus late views. Ratio effects for both magnitudes were observed starting in the P100. Moreover, a congruity effect emerged within the P100. That interactions were observed early in processing, at the same time that initial ratio effects occurred, suggests that number and cumulative area processes interacted when magnitude representations were being formed, prior to preparing a decision response. Our findings are consistent with an early interaction view of magnitude processing, in which number and cumulative area may rely on shared perceptual mechanisms.