Structural and functional connectivity of the inferior temporal numeral area.

A growing body of evidence suggests that in adults, there is a spatially consistent "inferior temporal numeral area" (ITNA) in the occipitotemporal cortex that appears to preferentially process Arabic digits relative to non-numerical symbols and objects. However, very little is known about why the ITNA is spatially segregated from regions that process other orthographic stimuli such as letters, and why it is spatially consistent across individuals. In the present study, we used diffusion-weighted imaging and functional magnetic resonance imaging to contrast structural and functional connectivity between left and right hemisphere ITNAs and a left hemisphere letter-preferring region. We found that the left ITNA had stronger structural and functional connectivity than the letter region to inferior parietal regions involved in numerical magnitude representation and arithmetic. Between hemispheres, the left ITNA showed stronger structural connectivity with the left inferior frontal gyrus (Broca's area), while the right ITNA showed stronger structural connectivity to the ipsilateral inferior parietal cortex and stronger functional coupling with the bilateral IPS. Based on their relative connectivity, our results suggest that the left ITNA may be more readily involved in mapping digits to verbal number representations, while the right ITNA may support the mapping of digits to quantity representations.

Probing the mechanisms underlying numerosity-to-numeral mappings and their relation to math competence.

Numerosity estimation performance (e.g., how accurate, consistent, or proportionally spaced (linear) numerosity-numeral mappings are) has previously been associated with math competence. However, the specific mechanisms that underlie such a relation is unknown. One possible mechanism is the mapping process between numerical sets and symbolic numbers (e.g., Arabic numerals). The current study examined two hypothesized mechanisms of numerosity-numeral mappings (item-based "associative" and holistic "structural" mapping) and their roles in the estimation-and-math relation. Specifically, mappings for small numbers (e.g., 1-10) are thought to be associative and resistant to calibration (e.g., feedback on accuracy of estimates), whereas holistic "structural" mapping for larger numbers (e.g., beyond 10) may be supported by flexibly aligning a numeral "response grid" (akin to a ruler) to an analog "mental number line" upon calibration. In 57 adults, we used pre- and post-calibration estimates to measure the range of continuous associative mappings among small numbers (e.g., a base range of associative mappings from 1 to 10), and obtained measures of math competence and delayed multiple-choice strategy reports. Consistent with previous research, uncalibrated estimation performance correlated with calculation competence, controlling for reading fluency and working memory. However, having a higher base range of associative mappings was not related to estimation performance or any math competence measures. Critically, discontinuity in calibration effects was typical at the individual level, which calls into question the nature of "holistic structural mapping". A parsimonious explanation to integrate previous and current findings is that estimation performance is likely optimized by dynamically constructing numerosity-numeral mappings through the use of multiple strategies from trial to trial.

The "Inferior Temporal Numeral Area" distinguishes numerals from other character categories during passive viewing: A representational similarity analysis.

A region in the posterior inferior temporal gyrus (pITG) is thought to be specialized for processing Arabic numerals, but fMRI studies that compared passive viewing of numerals to other character types (e.g., letters and novel characters) have not found evidence of numeral preference in the pITG. However, recent studies showed that the engagement of the pITG is modulated by attention and task contexts, suggesting that passive viewing paradigms may be ill-suited for examining numeral specialization in the pITG. It is possible, however, that even if the strengths of responses to different category types are similar, the distributed response patterns (i.e., neural representations) in a candidate numeral-preferring pITG region ("pITG-numerals") may reveal categorical distinctions, even during passive viewing. Using representational similarity analyses with three datasets that share the same task paradigm and stimulus sets (total N â€‹= â€‹88), we tested whether the neural representations of digits, letters, and novel characters in pITG-numerals were organized according to visual form and/or conceptual categories (e.g., familiar versus novel, numbers versus others). Small-scale frequentist and Bayesian meta-analyses of our dataset-specific findings revealed that the organization of neural representations in pITG-numerals is unlikely to be described by differences in abstract shape, but can be described by a categorical "digits versus letters" distinction, or even a "digits versus others" distinction (suggesting greater numeral sensitivity). Evidence of greater numeral sensitivity during passive viewing suggest that pITG-numerals is likely part of a neural pathway that has been developed for automatic processing of objects with potential numerical relevance. Given that numerals and letters do not differ categorically in terms of shape, categorical distinction in pITG-numerals during passive viewing must reflect ontogenetic differentiation of symbol set representations based on repeated usage of numbers and letters in differing task contexts.

The relation between 1st grade grey matter volume and 2nd grade math competence.

Mathematical and numerical competence is a critical foundation for individual success in modern society yet the neurobiological sources of individual differences in math competence are poorly understood. Neuroimaging research over the last decade suggests that neural mechanisms in the parietal lobe, particularly the intraparietal sulcus (IPS) are structurally aberrant in individuals with mathematical learning disabilities. However, whether those same brain regions underlie individual differences in math performance across the full range of math abilities is unknown. Furthermore, previous studies have been exclusively cross-sectional, making it unclear whether variations in the structure of the IPS are caused by or consequences of the development of math skills. The present study investigates the relation between grey matter volume across the whole brain and math competence longitudinally in a representative sample of 50 elementary school children. Results show that grey matter volume in the left IPS at the end of 1st grade relates to math competence a year later at the end of 2nd grade. Grey matter volume in this region did not change over that year, and was still correlated with math competence at the end of 2nd grade. These findings support the hypothesis that the IPS and its associated functions represent a critical foundation for the acquisition of mathematical competence.

Functional and representational differences between bilateral inferior temporal numeral areas.

The processing of numerals as visual objects is supported by an "Inferior Temporal Numeral Area" (ITNA) in the bilateral inferior temporal gyri (ITG). Extant findings suggest some degree of hemispheric asymmetry in how the bilateral ITNAs process numerals. Pollack and Price (2019) reported such a hemispheric asymmetry by which a region in the left ITG was sensitive to digits during a visual search for a digit among letters, and a homologous region in the right ITG that showed greater digit sensitivity in individuals with higher calculation skills. However, the ITG regions were localized with separate analyses without directly contrasting their digit sensitivities and relation to calculation skills. So, the extent of and reasons for these functional asymmetries remain unclear. Here we probe whether the functional and representational properties of the ITNAs are asymmetric by applying both univariate and multivariate region-of-interest analyses to Pollack and Price's (2019) data. Contrary to the implications of the original findings, digit sensitivity did not differ between ITNAs, and digit sensitivity in both left and right ITNAs was associated with calculation skills. Representational similarity analyses revealed that the overall representational geometries of digits in the ITNAs were also correlated, albeit weakly, but the representational contents of the ITNAs were largely inconclusive. Nonetheless, we found a right lateralization in engagement in alphanumeric categorization, and that the right ITNA showed greater discriminability between digits and letters. Greater right lateralization of digit sensitivity and digit discriminability in the left ITNA were also related to higher calculation skills. Our findings thus suggest that the ITNAs may not be functionally identical and should be directly contrasted in future work. Our study also highlights the importance of within-individual comparisons for understanding hemispheric asymmetries, and analyses of individual differences and multivariate features to uncover effects that would otherwise be obscured by averages.

Network topology of symbolic and nonsymbolic number comparison.

Studies of brain activity during number processing suggest symbolic and nonsymbolic numerical stimuli (e.g., Arabic digits and dot arrays) engage both shared and distinct neural mechanisms. However, the extent to which number format influences large-scale functional network organization is unknown. In this study, using 7 Tesla MRI, we adopted a network neuroscience approach to characterize the whole-brain functional architecture supporting symbolic and nonsymbolic number comparison in 33 adults. Results showed the degree of global modularity was similar for both formats. The symbolic format, however, elicited stronger community membership among auditory regions, whereas for nonsymbolic, stronger membership was observed within and between cingulo-opercular/salience network and basal ganglia communities. The right posterior inferior temporal gyrus, left intraparietal sulcus, and two regions in the right ventromedial occipital cortex demonstrated robust differences between formats in terms of their community membership, supporting prior findings that these areas are differentially engaged based on number format. Furthermore, a unified fronto-parietal/dorsal attention community in the nonsymbolic condition was fractionated into two components in the symbolic condition. Taken together, these results reveal a pattern of overlapping and distinct network architectures for symbolic and nonsymbolic number processing.

Shared Numerosity Representations Across Formats and Tasks Revealed with 7 Tesla fMRI: Decoding, Generalization, and Individual Differences in Behavior.

Debate continues on whether encoding of symbolic number is grounded in nonsymbolic numerical magnitudes. Nevertheless, fluency of perceiving both number formats, and translating between them, predicts math skills across the life span. Therefore, this study asked if numbers share cortical activation patterns across formats and tasks, and whether neural response to number predicts math-related behaviors. We analyzed patterns of neural activation using 7 Tesla functional magnetic resonance imaging in a sample of 39 healthy adults. Discrimination was successful between numerosities 2, 4, 6, and 8 dots and generalized to activation patterns of the same numerosities represented as Arabic digits in the bilateral parietal lobes and left inferior frontal gyrus (IFG) (and vice versa). This indicates that numerosity-specific neural resources are shared between formats. Generalization was also successful across tasks where participants either identified or compared numerosities in bilateral parietal lobes and IFG. Individual differences in decoding did not relate to performance on a number comparison task completed outside of the scanner, but generalization between formats and across tasks negatively related to math achievement in the parietal lobes. Together, these findings suggest that individual differences in representational specificity within format and task contexts relate to mathematical expertise.

Malleability of mappings between Arabic numerals and approximate quantities: Factors underlying individual differences and the relation to math.

Humans tend to be inaccurate and inconsistent when estimating a large number of objects. Furthermore, we modify our estimates when feedback or a reference array is provided, indicating that the mappings between perceived numerosity and their corresponding numerals are largely malleable in response to calibration. However, there is great variability in response to calibration across individuals. Using uncalibrated and calibrated numerosity estimation conditions, the current study explored the factors underlying individual differences in the extent and nature of the malleability of numerosity estimation performance as a result of calibration in a sample of 71 undergraduate students. We found that individual differences in performance were reliable across conditions, and participants' responses to calibration varied greatly. Participants who were less consistent or had more proportionally spaced (i.e., linear) estimates before calibration tended to shift the distributions of their estimates to a greater extent. Higher calculation competence also predicted an increase in how linear participants' estimates were after calibration. Moreover, the effect of calibration was not continuous across numerosities within participants. This suggests that the mechanisms underlying numeral-numerosity mappings may be less systematic than previously thought and likely depend on cognitive mechanisms beyond representation of numerosities. Taken together, the mappings between numerosities and numerical symbols may not be stable and direct, but transient and mediated by task-related (e.g., strategic) mechanisms. Rather than estimation skills being foundational for math competence, math competence may also influence estimation skills. Therefore, numerosity estimation tasks are not a pure measure of number representations.

Prospective relations between resting-state connectivity of parietal subdivisions and arithmetic competence.

The present study investigates the relation between resting-state functional connectivity (rsFC) of cytoarchitectonically defined subdivisions of the parietal cortex at the end of 1st grade and arithmetic performance at the end of 2nd grade. Results revealed a dissociable pattern of relations between rsFC and arithmetic competence among subdivisions of intraparietal sulcus (IPS) and angular gyrus (AG). rsFC between right hemisphere IPS subdivisions and contralateral IPS subdivisions positively correlated with arithmetic competence. In contrast, rsFC between the left hIP1 and the right medial temporal lobe, and rsFC between the left AG and left superior frontal gyrus, were negatively correlated with arithmetic competence. These results suggest that strong inter-hemispheric IPS connectivity is important for math development, reflecting either neurocognitive mechanisms specific to arithmetic processing, domain-general mechanisms that are particularly relevant to arithmetic competence, or structural 'cortical maturity'. Stronger connectivity between IPS, and AG, subdivisions and frontal and temporal cortices, however, appears to be negatively associated with math development, possibly reflecting the ability to disengage suboptimal problem-solving strategies during mathematical processing, or to flexibly reorient task-based networks. Importantly, the reported results pertain even when controlling for reading, spatial attention, and working memory, suggesting that the observed rsFC-behavior relations are specific to arithmetic competence.

The search for the number form area: A functional neuroimaging meta-analysis.

Recent studies report a putative "number form area" (NFA) in the inferior temporal gyrus (ITG) suggested to be specialized for Arabic numeral processing. However, a number of earlier studies report no such NFA. The reasons for such discrepancies across studies are unclear. To examine evidence for a convergent NFA across studies, we conducted two activation likelihood estimation meta-analyses on 31 and a subset of 20 neuroimaging studies that have contrasted digits with other meaningful symbols. Results suggest the potential existence of an NFA in the right ITG, in addition to a 'symbolic number processing network' comprising bilateral parietal regions, and right-lateralized superior and inferior frontal regions. Critically, convergent localization for the NFA was only evident when contrasts were appropriately controlled for task demands, and does not appear to depend on employing methods designed to overcome fMRI signal dropout in the ITG. Importantly, only five studies had foci within the identified ITG NFA cluster boundary, indicating that more empirical evidence is necessary to determine the true functional specialization and regional specificity of the putative NFA.

Eye-movement patterns during nonsymbolic and symbolic numerical magnitude comparison and their relation to math calculation skills.

A growing body of research suggests that the processing of nonsymbolic (e.g. sets of dots) and symbolic (e.g. Arabic digits) numerical magnitudes serves as a foundation for the development of math competence. Performance on magnitude comparison tasks is thought to reflect the precision of a shared cognitive representation, as evidence by the presence of a numerical ratio effect for both formats. However, little is known regarding how visuo-perceptual processes are related to the numerical ratio effect, whether they are shared across numerical formats, and whether they relate to math competence independently of performance outcomes. The present study investigates these questions in a sample of typically developing adults. Our results reveal a pattern of associations between eye-movement measures, but not their ratio effects, across formats. This suggests that ratio-specific visuo-perceptual processing during magnitude processing is different across nonsymbolic and symbolic formats. Furthermore, eye movements are related to math performance only during symbolic comparison, supporting a growing body of literature suggesting symbolic number processing is more strongly related to math outcomes than nonsymbolic magnitude processing. Finally, eye-movement patterns, specifically fixation dwell time, continue to be negatively related to math performance after controlling for task performance (i.e. error rate and reaction time) and domain general cognitive abilities (IQ), suggesting that fluent visual processing of Arabic digits plays a unique and important role in linking symbolic number processing to formal math abilities.