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.

Top-down determinants of the numerosity-time interaction.

Previous studies have reported that larger visual stimuli are perceived as lasting longer than smaller ones. However, this effect disappears when participants provide a qualitative judgment, by stating whether two stimuli have the "same or different" duration, instead of providing an explicit quantitative judgment (which stimulus lasts longer). Here, we extended these observations to the interaction between the numerosity of visual stimuli, i.e. clouds of dots, and their duration. With "longer vs shorter" responses, participants judged larger numerosities as lasting longer than smaller ones, both when the responses were related to the order (Experiment 1) or color (Experiment 4) of stimuli. In contrast, no similar effect was found with "same vs different" responses (Experiment 2) and in a time motor reproduction task (Experiment 3). The numerosity-time interference in Experiment 1 and Experiment 4 was not due to task difficulty, as sensory precision was equivalent to that of Experiment 2. We conclude that in humans the functional interaction between numerosity and time is not guided, in the main, by a shared bottom-up mechanism of magnitude coding. Rather, high-level and top-down processes involved in decision-making and guided by the use of "magnitude-related" response codes play a crucial role in triggering interference among different magnitude domains.

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.

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.

Similar effect of running on visual and auditory time perception in the ranges of milliseconds and seconds.

Introduction: The ability to accurately encode events' duration is of critical importance for almost all everyday activities, yet numerous factors have been reported to robustly distort time perception. One of these is physical activity (i.e., running, walking) but, partly due to the variety of methodologies employed, a full comprehension of the role of exercise on the encoding of time has still to be achieved. Methods: Here we tackle the issue with a multifaceted approach by measuring the effect of vigorous running with a time generalization task for visual and auditory stimuli in the range of milliseconds (0.2-0.8 s) as well as seconds (1-4 s). At baseline, participants performed both the encoding and decoding at rest while in the experimental conditions the decoding was performed while running. Results: Our results indicate that physical activity in both duration ranges (sub-second and seconds) was expanded during running regardless of the sensory modality used to present the stimuli. Despite this generalized effect of running on perceived duration, we found evidence for the existence of independent timing mechanisms: (1) the perceptual biases induced by running in the two temporal regimes were uncorrelated, (2) sensory precision levels (Weber fraction) were higher for stimuli in the seconds range, (3) sensory precision levels were higher for auditory than for visual stimuli, but only within the sub-second range. Discussion: Overall, our results support previous findings suggesting (at least partially) separate timing mechanisms for short/long durations and for visual and auditory stimuli. However, they also indicate that physical activity affects all these temporal modules, suggesting a generalized interaction-via generalized and shared resources-between the motor system and the brain time mechanisms.

Visual P2p component responds to perceived numerosity.

Numerosity perception is a key ability for human and non-human species, probably mediated by dedicated brain mechanisms. Electrophysiological studies revealed the existence of both early and mid-latency components of the Electrophysiological (EEG) signal sensitive to numerosity changes. However, it is still unknown whether these components respond to physical or perceived variation in numerical attributes. We here tackled this point by recording electrophysiological signal while participants performed a numerosity adaptation task, a robust psychophysical method yielding changes in perceived numerosity judgments despite physical numerosity invariance. Behavioral measures confirmed that the test stimulus was consistently underestimated when presented after a high numerous adaptor while perceived as veridical when presented after a neutral adaptor. Congruently, EEG results revealed a potential at around 200 ms (P2p) which was reduced when the test stimulus was presented after the high numerous adaptor. This result was much prominent over the left posterior cluster of electrodes and correlated significantly with the amount of adaptation. No earlier modulations were retrievable when changes in numerosity were illusory while both early and mid-latency modulations occurred for physical changes. Taken together, our results reveal that mid-latency P2p mainly reflects perceived changes in numerical attributes, while earlier components are likely to be bounded to the physical characteristics of the stimuli. These results suggest that short-term plastic mechanisms induced by numerosity adaptation may involve a relatively late processing stage of the visual hierarchy likely engaging cortical areas beyond the primary visual cortex. Furthermore, these results also indicate mid-latency electrophysiological correlates as a signature of the internal representation of numerical information.

Math difficulties in attention deficit hyperactivity disorder do not originate from the visual number sense.

There is ample evidence from literature and clinical practice indicating mathematical difficulties in individuals with ADHD, even when there is no concomitant diagnosis of developmental dyscalculia. What factors underlie these difficulties is still an open question. Research on dyscalculia and neurotypical development suggests visual perception of numerosity (the number sense) as a building block for math learning. Participants with lower numerosity estimation thresholds (higher precision) are often those with higher math capabilities. Strangely, the role of numerosity perception in math skills in ADHD has been neglected, leaving open the question whether math difficulties in ADHD also originate from a deficitary visual number sense. In the current study we psychophysically measured numerosity thresholds and accuracy in a sample of children/adolescents with ADHD, but not concomitant dyscalculia (N = 20, 8-16 years). Math abilities were also measured by tasks indexing different mathematical competences. Numerosity performance and math scores were then compared to those obtained from an age-matched control group (N = 20). Bayesian statistics indicated no difference between ADHD and controls on numerosity perception, despite many of the symbolic math tasks being impaired in participants with ADHD. Moreover, the math deficits showed by the group with ADHD remained substantial even when numerosity thresholds were statistically regressed out. Overall, these results indicate that math difficulties in ADHD are unlikely to originate from an impaired visual number sense.

Symmetry as a grouping cue for numerosity perception.

To estimate the number of objects in an image, each element needs to be segregated as a single unit. Several principles guide the process of element identification, one of the strongest being symmetry. In the current study, we investigated how symmetry affects the ability to rapidly estimate the number of objects (numerosity). Participants judged the numerosity of asymmetric or symmetric arrays of various numerosities. The results show that the numerosity of symmetrical arrays was significantly underestimated at low numerosities, but the effect was greatly reduced at higher numerosities. Adding an additional axis of symmetry (double symmetry) further reduced perceived numerosity. The magnitude of the symmetry-driven underestimation was inversely correlated with autistic personality traits, consistent with previous work associating autistic traits with perceptual grouping. Overall, these results support the idea that perceived numerosity relies on object segmentation and grouping cues, with symmetry playing a key role.

Mathematics and Numerosity but Not Visuo-Spatial Working Memory Correlate with Mathematical Anxiety in Adults.

Many individuals, when faced with mathematical tasks or situations requiring arithmetic skills, experience exaggerated levels of anxiety. Mathematical anxiety (MA), in addition to causing discomfort, can lead to avoidance behaviors and then to underachievement. However, the factors inducing MA and how MA deploys its detrimental effects are still largely debated. There is evidence suggesting that MA affects working memory capacity by further diminishing its limited processing resources. An alternative account postulates that MA originates from a coarse early numerical cognition capacity, the perception of numerosity. In the current study, we measured MA, math abilities, numerosity perception and visuo-spatial working memory (VSWM) in a sample of neurotypical adults. Correlational analyses confirmed previous studies showing that high MA was associated with lower math scores and worse numerosity estimation precision. Conversely, MA turned out to be unrelated to VSWM capacities. Finally, partial correlations revealed that MA fully accounted for the relationship between numerosity estimation precision and math abilities, suggesting a key role for MA as a mediating factor between these two domains.

Numerosity perception is tuned to salient environmental features.

Numerosity perception is a key ability to guide behavior. However, current models propose that number units encode an abstract representation of numerosity regardless of the non-numerical attributes of the stimuli, suggesting rather coarse environmental tuning. Here we investigated whether numerosity systems spontaneously adapt to all visible items, or to subsets segregated by salient attributes such as color or pitch. We measured perceived numerosity after participants adapted to highly numerous stimuli with color either matched to or different from the test. Matched colors caused a 25% underestimation of numerosity, while different colors had virtually no effect. This was true both for physically different colors, and for the same colors perceived as different, via a color-assimilation illusion. A similar result occurred in the acoustic domain, where adaptation magnitude was halved when the adaptor and test differed in pitch. Taken together, our results support the idea that numerosity perception is selectively tuned to salient environmental attributes.

Unimpaired groupitizing in children and adolescents with dyscalculia.

When asked to estimate the number of items in the visual field, neurotypical adults are more precise and rapid if the items are clustered into subgroups compared to when they are randomly distributed. It has been suggested that this phenomenon, termed "groupitizing", relies on the recruitment of arithmetical calculation strategies and subitizing. Here the role of arithmetical skills in groupitizing was investigated by measuring the groupitizing effect (or advantage) in a sample of children and adolescents with and without math learning disability (dyscalculia). The results showed that when items were grouped, both groups of participants showed a similar advantage on sensory precision and response time in numerosity estimates. Correlational analyses confirmed a lack of covariation between groupitizing advantage and math scores. Bayesian statistics on sensory precision sustained the frequentist analyses providing decisive evidence in favor of no groups difference on groupitizing advantage magnitude (LBF = - 0.44) and no correlation with math scores (LBF = - 0.57). The results on response times, although less decisive, were again in favor of the null hypothesis. Overall, the results suggest that the link between groupitizing and mathematical abilities cannot be taken for granted, calling for further investigations on the factors underlying this perceptual phenomenon.

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.

Visual Duration but Not Numerosity Is Distorted While Running.

There is increasing evidence that action and perception interact in the processing of magnitudes such as duration and numerosity. Sustained physical exercise (such as running or cycling) increases the apparent duration of visual stimuli presented during the activity. However, the effect of exercise on numerosity perception has not yet been investigated. Here, we asked participants to make either a temporal or a numerical judgment by comparing the duration or numerosity of standard stimuli displayed at rest with those presented while running. The results support previous reports in showing that physical activity significantly expands perceived duration; however, it had no effect on perceived numerosity. Furthermore, the distortions of the perceived durations vanished soon after the running session, making it unlikely that physiological factors such as heart rate underlie the temporal distortion. Taken together, these results suggest a domain-selective influence of the motor system on the perception of time, rather than a general effect on magnitude.

Auditory time thresholds in the range of milliseconds but not seconds are impaired in ADHD.

The literature on time perception in individuals with ADHD is extensive but inconsistent, probably reflecting the use of different tasks and performances indexes. A sample of 40 children/adolescents (20 with ADHD, 20 neurotypical) was engaged in two identical psychophysical tasks measuring auditory time thresholds in the milliseconds (0.25-1 s) and seconds (0.75-3 s) ranges. Results showed a severe impairment in ADHD for milliseconds thresholds (Log10BF = 1.9). The deficit remained strong even when non-verbal IQ was regressed out and correlation with age suggests a developmental delay. In the seconds range, thresholds were indistinguishable between the two groups (Log10BF = - 0.5) and not correlated with milliseconds thresholds. Our results largely confirm previous evidence suggesting partially separate mechanisms for time perception in the ranges of milliseconds and seconds. Moreover, since the evidence suggests that time perception of milliseconds stimuli might load relatively less on cognitive control and working memory, compared to longer durations, the current results are consistent with a pure timing deficit in individuals with ADHD.

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.

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.