What pupil size can and cannot tell about math anxiety.

Math Anxiety (MA) consists of excessive fear and worry about math-related situations. It represents a major barrier to numerical competence and the pursuit of STEM careers. Yet, we currently do not dispose of many tools that can capture its multifaceted nature, e.g. moving beyond the exclusive reliance on self-reports and meta-cognition. Here we sought to probe Pupil Size (PS) as a viable tool in the study of MA by administering arithmetic problems to university students in the humanities (N = 70) with various levels of MA. We found that arithmetic competence and performance are indeed negatively associated with MA, and this is accurately tracked by PS. When performance is accounted for, MA does not further modulate PS (before, during, or after calculation). However, the latency of PS peak dilation can add a significant contribution to predicting MA scores, indicating that high MA may be accompanied by more prolonged cognitive effort. Results show that MA and mathematical competence may be too crystalized in young university students to be discernible. We therefore call for early educational interventions to tackle and mitigate this dysfunctional association early on.

The role of motor effort on the sensorimotor number system.

The integration of numerical information with motor processes has emerged as a fascinating area of investigation in both animal and human cognition. The interest in a sensorimotor number system has recently generated neurophysiological and psychophysical evidence which combine to highlight the importance of motor functions in the encoding of numerical information. Nevertheless, several key questions remain, such as the influence of non-numerical motor parameters over numerical perception. Here we tested the role of physical effort, a parameter positively correlated with the number of actions, in modulating the link between hand-actions and visual numerosity perception. Effort was manipulated during sensorimotor adaptation as well as during a new actions-estimation paradigm. The results of Experiment 1 shows that physical effort in the absence of actions (passive effort) is not sufficient to activate the sensorimotor number system, indicating that self-produced actions are instead necessary. Further experiments demonstrated that effort is marginally integrated during motor adaptation (Experiment 2) but discarded when estimating the number of self-produced hand actions (Experiment 3). Overall, the results indicate that the sensorimotor number system is largely fed by the number of discrete actions rather than the amount of effort but also indicates that effort (under specific circumstances) might be integrated. These findings provide novel insights into the sensorimotor numerical integration, paving the way for future investigations, such as on its functional role.

Groupitizing modifies neural coding of numerosity.

Numerical estimation of arrays of objects is faster and more accurate when items can be clustered into groups, a phenomenon termed "groupitizing." Grouping can facilitate segregation into subitizable "chunks," each easily estimated, then summed. The current study investigates whether spatial grouping of arrays drives specific neural responses during numerical estimation, reflecting strategies such as exact calculation and fact retrieval. Fourteen adults were scanned with fMRI while estimating either the numerosity or shape of arrays of items, either randomly distributed or spatially grouped. Numerosity estimation of both classes of stimuli elicited common activation of a right lateralized frontoparietal network. Grouped stimuli additionally recruited regions in the left hemisphere and bilaterally in the angular gyrus. Multivariate pattern analysis showed that classifiers trained with the pattern of neural activations read out from parietal regions, but not from the primary visual areas, can decode different numerosities both within and across spatial arrangements. The behavioral numerical acuity correlated with the decoding performance of the parietal but not with occipital regions. Overall, this experiment suggests that the estimation of grouped stimuli relies on the approximate number system for numerosity estimation, but additionally recruits regions involved in calculation.

Uncertainty and Prior Assumptions, Rather Than Innate Logarithmic Encoding, Explain Nonlinear Number-to-Space Mapping.

Mapping number to space is natural and spontaneous but often nonveridical, showing a clear compressive nonlinearity that is thought to reflect intrinsic logarithmic encoding of numerical values. We asked 78 adult participants to map dot arrays onto a number line across nine trials. Combining participant data, we confirmed that on the first trial, mapping was heavily compressed along the number line, but it became more linear across trials. Responses were well described by logarithmic compression but also by a parameter-free Bayesian model of central tendency, which quantitatively predicted the relationship between nonlinearity and number acuity. To experimentally test the Bayesian hypothesis, we asked 90 new participants to complete a color-line task in which they mapped noise-perturbed color patches to a "color line." When there was more noise at the high end of the color line, the mapping was logarithmic, but it became exponential with noise at the low end. We conclude that the nonlinearity of both number and color mapping reflects contextual Bayesian inference processes rather than intrinsic logarithmic encoding.

Normal Retinotopy in Primary Visual Cortex in a Congenital Complete Unilateral Lesion of Lateral Geniculate Nucleus in Human: A Case Study.

Impairment of the geniculostriate pathway results in scotomas in the corresponding part of the visual field. Here, we present a case of patient IB with left eye microphthalmia and with lesions in most of the left geniculostriate pathway, including the Lateral Geniculate Nucleus (LGN). Despite the severe lesions, the patient has a very narrow scotoma in the peripheral part of the lower-right-hemifield only (beyond 15° of eccentricity) and complete visual field representation in the primary visual cortex. Population receptive field mapping (pRF) of the patient's visual field reveals orderly eccentricity maps together with contralateral activation in both hemispheres. With diffusion tractography, we revealed connections between superior colliculus (SC) and cortical structures in the hemisphere affected by the lesions, which could mediate the retinotopic reorganization at the cortical level. Our results indicate an astonishing case for the flexibility of the developing retinotopic maps where the contralateral thalamus receives fibers from both the nasal and temporal retinae.

Numerosity adaptation partly depends on the allocation of implicit numerosity-contingent visuo-spatial attention.

Like other perceptual attributes, numerosity is susceptible to adaptation. Nevertheless, it has never been fully investigated whether adaptation to numerosity is fully perceptual in nature or if it stems from the mixed influence of perception and attention. In the present work, we addressed this point throughout three separate experiments aiming at investigating the potential role played by visuo-spatial attentional mechanisms in shaping numerosity perception and adaptation. In Experiments 1 and 2, we showed that the magnitude of numerosity adaptation can be strongly influenced by the distribution of numerosity-contingent visuo-spatial attentional resources during the adaptation period. Results from Experiment 1 revealed a robust reduction of adaptation magnitude whenever a second numerical stimulus was presented in a diametrically opposite location from that of the adaptor, despite this second adapter being neutral as matched in numerosity with the following stimulus displayed in that location. In Experiment 2, we showed that this reduction in adaptation did not occur in cases where the second stimulus was not numerical, suggesting that attentional resources specifically related to numerosity information accounts for the results of Experiment 1. Finally, in Experiment 3, we showed that uninformative visuo-spatial cues shape numerosity discrimination judgments both at baseline and during adaptation. Taken together, our results seem to indicate that visuo-spatial attention plays a relevant role in numerosity perception and that adaptation to numerosity is actively influenced by this cognitive process.

Implicit visuospatial attention shapes numerosity adaptation and perception.

The perception of numerical quantities is susceptible to adaptation: after inspecting a numerous dot array for a few seconds a subsequent dot array is grossly underestimated. In a recent work we showed that the mere appearance of an additional numerically neutral stimulus significantly reduces the adaptation magnitude. Here we demonstrate that this reduction is likely due to a numerosity underestimation of the adaptor caused by a change of numerosity-related attentional resources deployed on the adapting stimulus. In Experiment 1 we replicated previous findings revealing a robust reduction of numerosity adaptation when an additional adaptor (even if neutral) was displayed. In Experiment 2 we used the method of magnitude estimation to demonstrate that numerosity is underestimated whenever a second task-irrelevant numerical stimulus appears on screen. Furthermore we demonstrated that the same experimental manipulations were not effective in modulating orientation adaptation magnitude as well as orientation estimation accuracy. Our results support the hypothesis of a tight relationship between numerosity perception and implicit visuospatial attention and corroborate the notion that numerosity adaptation depends on perceived rather than physical numerosity. However the lack of an effect of visuospatial attentional deployment for orientation perception suggests that attention might differently shape adaptation aftereffects for different features along the visual hierarchy.

Numbers in action.

To understand the number sense, we need to understand its function. We argue that numerosity estimation is fundamental not only for perception, but also preparation and control of action. We outline experiments that link numerosity estimation with action, pointing to a generalized numerosity system that serves both perception and action preparation.

Simultaneous and sequential subitizing are separate systems, and neither predicts math abilities.

Small quantities of visual objects can be rapidly estimated without error, a phenomenon known as subitizing. Larger quantities can also be rapidly estimated, but with error, and the error rate predicts math abilities. This study addressed two issues: (a) whether subitizing generalizes over modalities and stimulus formats and (b) whether subitizing correlates with math abilities. We measured subitizing limits in primary school children and adults for visual and auditory stimuli presented either sequentially (sequences of flashes or sounds) or simultaneously (visual presentations, dot arrays). The results show that (a) subitizing limits for adults were one item larger than those for primary school children across all conditions; (b) subitizing for simultaneous visual stimuli (dots) was better than that for sequential stimuli; (c) subitizing limits for dots do not correlate with subitizing limits for either flashes or sounds; (d) subitizing of sequences of flashes and subitizing of sequences of sounds are strongly correlated with each other in children; and (e) regardless of stimuli sensory modality and format, subitizing limits do not correlate with mental calculation or digit magnitude knowledge proficiency. These results suggest that although children can subitize sequential numerosity, simultaneous and temporal subitizing may be subserved by separate systems. Furthermore, subitizing does not seem to be related to numerical abilities.

Different reaction-times for subitizing, estimation, and texture.

Humans can estimate and encode numerosity over a large range, from very few items to several hundreds. Two distinct mechanisms have been proposed: subitizing, for numbers up to four and estimation for larger numerosities. We have recently extended this idea by suggesting that for very densely packed arrays, when items are less segregable, a third "texture" mechanism comes into play. In this study, we provide further evidence for the existence of a third regime for numerosity. Reaction times were very low in the subitizing range, rising rapidly for numerosities greater than four. However, for tightly packed displays of very high numerosities, reaction times became faster. These results reinforce the idea of three regimes in the processing of numerosity, subitizing, estimation, and texture.

Compressive mapping of number to space reflects dynamic encoding mechanisms, not static logarithmic transform.

The mapping of number onto space is fundamental to measurement and mathematics. However, the mapping of young children, unschooled adults, and adults under attentional load shows strong compressive nonlinearities, thought to reflect intrinsic logarithmic encoding mechanisms, which are later "linearized" by education. Here we advance and test an alternative explanation: that the nonlinearity results from adaptive mechanisms incorporating the statistics of recent stimuli. This theory predicts that the response to the current trial should depend on the magnitude of the previous trial, whereas a static logarithmic nonlinearity predicts trialwise independence. We found a strong and highly significant relationship between numberline mapping of the current trial and the magnitude of the previous trial, in both adults and school children, with the current response influenced by up to 15% of the previous trial value. The dependency is sufficient to account for the shape of the numberline, without requiring logarithmic transform. We show that this dynamic strategy results in a reduction of reproduction error, and hence improvement in accuracy.

Mechanisms for perception of numerosity or texture-density are governed by crowding-like effects.

We have recently provided evidence that the perception of number and texture density is mediated by two independent mechanisms: numerosity mechanisms at relatively low numbers, obeying Weber's law, and texture-density mechanisms at higher numerosities, following a square root law. In this study we investigated whether the switch between the two mechanisms depends on the capacity to segregate individual dots, and therefore follows similar laws to those governing visual crowding. We measured numerosity discrimination for a wide range of numerosities at three eccentricities. We found that the point where the numerosity regime (Weber's law) gave way to the density regime (square root law) depended on eccentricity. In central vision, the regime changed at 2.3 dots/°2, while at 15° eccentricity, it changed at 0.5 dots/°2, three times less dense. As a consequence, thresholds for low numerosities increased with eccentricity, while at higher numerosities thresholds remained constant. We further showed that like crowding, the regime change was independent of dot size, depending on distance between dot centers, not distance between dot edges or ink coverage. Performance was not affected by stimulus contrast or blur, indicating that the transition does not depend on low-level stimulus properties. Our results reinforce the notion that numerosity and texture are mediated by two distinct processes, depending on whether the individual elements are perceptually segregable. Which mechanism is engaged follows laws that determine crowding.

Separate mechanisms for perception of numerosity and density.

Despite the existence of much evidence for a number sense in humans, several researchers have questioned whether number is sensed directly or derived indirectly from texture density. Here, we provide clear evidence that numerosity and density judgments are subserved by distinct mechanisms with different psychophysical characteristics. We measured sensitivity for numerosity discrimination over a wide range of numerosities: For low densities (less than 0.25 dots/deg(2)), thresholds increased directly with numerosity, following Weber's law; for higher densities, thresholds increased with the square root of texture density, a steady decrease in the Weber fraction. The existence of two different psychophysical systems is inconsistent with a model in which number is derived indirectly from noisy estimates of density and area; rather, it points to the existence of separate mechanisms for estimating density and number. These results provide strong confirmation for the existence of neural mechanisms that sense number directly, rather than indirectly from texture density.

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.

Auditory time perception impairment in children with developmental dyscalculia.

Developmental dyscalculia (DD) is a specific learning disability which prevents children from acquiring adequate numerical and arithmetical competences. We investigated whether difficulties in children with DD spread beyond the numerical domain and impact also their ability to perceive time. A group of 37 children/adolescent with and without DD were tested with an auditory categorization task measuring time perception thresholds in the sub-second (0.25-1 s) and supra-second (0.75-3 s) ranges. Results showed that auditory time perception was strongly impaired in children with DD at both time scales. The impairment remained even when age, non-verbal reasoning, and gender were regressed out. Overall, our results show that the difficulties of DD can affect magnitudes other than numerical and contribute to the increasing evidence that frames dyscalculia as a disorder affecting multiple neurocognitive and perceptual systems.

Neurocognitive Assessment of Mathematics-Related Capacities in Neurosurgical Patients.

A precise neuropsychological assessment is of the utmost importance for neurosurgical patients undergoing the surgical excision of cerebral lesions. The assessment of mathematical abilities is usually limited to arithmetical operations while other fundamental visuo-spatial aspects closely linked to mathematics proficiency, such as the perception of numerical quantities and geometrical reasoning, are completely neglected. We evaluated these abilities with two objective and reproducible psychophysical tests, measuring numerosity perception and non-symbolic geometry, respectively. We tested sixteen neuro-oncological patients before the operation and six after the operation with classical neuropsychological tests and with two psychophysical tests. The scores of the classical neuropsychological tests were very heterogeneous, possibly due to the distinct location and histology of the tumors that might have spared (or not) brain areas subserving these abilities or allowed for plastic reorganization. Performance in the two non-symbolic tests reflected, on average, the presumed functional role of the lesioned areas, with participants with parietal and frontal lesions performing worse on these tests than patients with occipital and temporal lesions. Single-case analyses not only revealed some interesting exceptions to the group-level results (e.g., patients with parietal lesions performing well in the numerosity test), but also indicated that performance in the two tests was independent of non-verbal reasoning and visuo-spatial working memory. Our results highlight the importance of assessing non-symbolic numerical and geometrical abilities to complement typical neuropsychological batteries. However, they also suggest an avoidance of reliance on an excessively rigid localizationist approach when evaluating the neuropsychological profile of oncological patients.

Visual sustained attention and numerosity sensitivity correlate with math achievement in children.

In this study, we investigated in school-age children the relationship among mathematical performance, the perception of numerosity (discrimination and mapping to number line), and sustained visual attention. The results (on 68 children between 8 and 11 years of age) show that attention and numerosity perception predict math scores but not reading performance. Even after controlling for several variables, including age, gender, nonverbal IQ, and reading accuracy, attention remained correlated with math skills and numerosity discrimination. These findings support previous reports showing the interrelationship between visual attention and both numerosity perception and math performance. It also suggests that attentional deficits may be implicated in disturbances such as developmental dyscalculia.

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.

The symmetry-induced numerosity illusion depends on visual attention.

Symmetry is an important and strong cue we rely on to organize the visual world. Although it is at the basis of objects segmentation in a visual scene, it can sometimes bias our perception. When asked to discriminate numerical quantities between symmetric and asymmetric arrays, individuals tend to underestimate the number of items in the symmetric stimuli. The reason for this underestimation is currently unknown. In this study we investigated whether the symmetry-induced numerosity underestimation depends on perceptual grouping mechanisms by depriving attentional resources. Twenty-six adults judged the numerosity of dot arrays arranged symmetrically or randomly, while ignoring a visual distractor (single task) or while simultaneously judging its color and orientation (dual-task). Diverting attention to the concurrent color-orientation conjunction task halved the symmetry-induced numerosity underestimation. Taken together these results showed that the bias in numerosity perception of symmetric arrays depends-at least partially-on attentional resources and suggested that it might originate from the recruitment of attentional dependent incremental grouping mechanisms.