Orthographic dependency in the neural correlates of reading: evidence from audiovisual integration in English readers.

Reading skills are indispensible in modern technological societies. In transparent alphabetic orthographies, such as Dutch, reading skills build on associations between letters and speech sounds (LS pairs). Previously, we showed that the superior temporal cortex (STC) of Dutch readers is sensitive to the congruency of LS pairs. Here, we used functional magnetic resonance imaging to investigate whether a similar congruency sensitivity exists in STC of readers of the more opaque English orthography, where the relation among LS pairs is less reliable. Eighteen subjects passively perceived congruent and incongruent audiovisual pairs of different levels of transparency in English: letters and speech sounds (LS; irregular), letters and letter names (LN; fairly transparent), and numerals and number names (NN; transparent). In STC, we found congruency effects for NN and LN, but no effects in the predicted direction (congruent > incongruent) for LS pairs. These findings contrast with previous results obtained from Dutch readers. These data indicate that, through education, the STC becomes tuned to the congruency of transparent audiovisual pairs, but suggests a different neural processing of irregular mappings. The orthographic dependency of LS integration underscores cross-linguistic differences in the neural basis of reading and potentially has important implications for dyslexia interventions across languages.

Semantic and perceptual processing of number symbols: evidence from a cross-linguistic fMRI adaptation study.

The ability to process the numerical magnitude of sets of items has been characterized in many animal species. Neuroimaging data have associated this ability to represent nonsymbolic numerical magnitudes (e.g., arrays of dots) with activity in the bilateral parietal lobes. Yet the quantitative abilities of humans are not limited to processing the numerical magnitude of nonsymbolic sets. Humans have used this quantitative sense as the foundation for symbolic systems for the representation of numerical magnitude. Although numerical symbol use is widespread in human cultures, the brain regions involved in processing of numerical symbols are just beginning to be understood. Here, we investigated the brain regions underlying the semantic and perceptual processing of numerical symbols. Specifically, we used an fMRI adaptation paradigm to examine the neural response to Hindu-Arabic numerals and Chinese numerical ideographs in a group of Chinese readers who could read both symbol types and a control group who could read only the numerals. Across groups, the Hindu-Arabic numerals exhibited ratio-dependent modulation in the left IPS. In contrast, numerical ideographs were associated with activation in the right IPS, exclusively in the Chinese readers. Furthermore, processing of the visual similarity of both digits and ideographs was associated with activation of the left fusiform gyrus. Using culture as an independent variable, we provide clear evidence for differences in the brain regions associated with the semantic and perceptual processing of numerical symbols. Additionally, we reveal a striking difference in the laterality of parietal activation between the semantic processing of the two symbols types.

Effects of problem size and arithmetic operation on brain activation during calculation in children with varying levels of arithmetical fluency.

Most studies on mathematics learning in the field of educational neuroscience have focused on the neural correlates of very elementary numerical processing skills in children. Little is known about more complex mathematical skills that are formally taught in school, such as arithmetic. Using functional magnetic resonance imaging, the present study investigated how brain activation during single-digit addition and subtraction is modulated by problem size and arithmetic operation in 28 children aged 10-12 years with different levels of arithmetical fluency. Commensurate with adult data, large problems and subtractions activated a fronto-parietal network, including the intraparietal sulci, the latter of which indicates the influence of quantity-based processes during procedural strategy execution. Different from adults, the present findings revealed that particularly the left hippocampus was active during the solution of those problems that are expected to be solved by means of fact retrieval (i.e. small problems and addition), suggesting a specific role of the hippocampus in the early stages of learning arithmetic facts. Children with low levels of arithmetical fluency showed higher activation in the right intraparietal sulcus during the solution of problems with a relatively small problem size, indicating that they continued to rely to a greater extent on quantity-based strategies on those problems that the children with relatively higher arithmetical fluency already retrieved from memory. This might represent a neural correlate of fact retrieval impairments in children with mathematical difficulties.

Linking brain-wide multivoxel activation patterns to behaviour: Examples from language and math.

A key goal of cognitive neuroscience is to find simple and direct connections between brain and behaviour. However, fMRI analysis typically involves choices between many possible options, with each choice potentially biasing any brain-behaviour correlations that emerge. Standard methods of fMRI analysis assess each voxel individually, but then face the problem of selection bias when combining those voxels into a region-of-interest, or ROI. Multivariate pattern-based fMRI analysis methods use classifiers to analyse multiple voxels together, but can also introduce selection bias via data-reduction steps as feature selection of voxels, pre-selecting activated regions, or principal components analysis. We show here that strong brain-behaviour links can be revealed without any voxel selection or data reduction, using just plain linear regression as a classifier applied to the whole brain at once, i.e. treating each entire brain volume as a single multi-voxel pattern. The brain-behaviour correlations emerged despite the fact that the classifier was not provided with any information at all about subjects' behaviour, but instead was given only the neural data and its condition-labels. Surprisingly, more powerful classifiers such as a linear SVM and regularised logistic regression produce very similar results. We discuss some possible reasons why the very simple brain-wide linear regression model is able to find correlations with behaviour that are as strong as those obtained on the one hand from a specific ROI and on the other hand from more complex classifiers. In a manner which is unencumbered by arbitrary choices, our approach offers a method for investigating connections between brain and behaviour which is simple, rigorous and direct.

Developmental specialization in the right intraparietal sulcus for the abstract representation of numerical magnitude.

Because number is an abstract quality of a set, the way in which a number is externally represented does not change its quantitative meaning. In this study, we examined the development of the brain regions that support format-independent representation of numerical magnitude. We asked children and adults to perform both symbolic (Hindu-Arabic numerals) and nonsymbolic (arrays of squares) numerical comparison tasks as well as two control tasks while their brains were scanned using fMRI. In a preliminary analysis, we calculated the conjunction between symbolic and nonsymbolic numerical comparison. We then examined in which brain regions this conjunction differed between children and adults. This analysis revealed a large network of visual and parietal regions that showed greater activation in adults relative to children. In our primary analysis, we examined age-related differences in the conjunction of symbolic and nonsymbolic comparison after subtracting the control tasks. This analysis revealed a much more limited set of regions including the right inferior parietal lobe near the intraparietal sulcus. In addition to showing increased activation to both symbolic and nonsymbolic magnitudes over and above activation related to response selection, this region showed age-related differences in the distance effect. Our findings demonstrate that the format-independent representation of numerical magnitude in the right inferior parietal lobe is the product of developmental processes of cortical specialization and highlight the importance of using appropriate control tasks when conducting developmental neuroimaging studies.

Common and segregated neural pathways for the processing of symbolic and nonsymbolic numerical magnitude: an fMRI study.

Numbers are everywhere in modern life. Looking out a window, one might see both symbolic numbers, like the numerals on a thermometer, and nonsymbolic quantities, such as the number of chickadees at a bird feeder. Although differences between symbolic and nonsymbolic numbers appear very salient, most research on numerical cognition has focused on similarities rather than differences between numerical stimulus formats. Thus, little is known about differences in the processing of symbolic and nonsymbolic numerical magnitudes. A recent computational model proposed that symbolic and nonsymbolic quantities undergo distinct encoding processes which then converge on a common neural representation of numerical magnitude (Verguts, T., Fias, W., 2004. Representation of number in animals and humans: a neural model. J. Cogn. Neurosci. 16 (9), 1493-1504.). Moreover, this model predicted that discrete brain regions underlie these encoding processes. Using functional magnetic resonance imaging, the present study tested the predictions of this model by examining the functional neuroanatomy of symbolic and nonsymbolic number processing. Nineteen adults compared the relative numerical magnitude of symbolic and nonsymbolic stimuli. An initial conjunction analysis revealed the right inferior parietal lobule to be significantly active in both symbolic and nonsymbolic numerical comparison. A contrast of the activation associated with symbolic and nonsymbolic stimuli revealed that both the left angular and superior temporal gyri were more activated for symbolic compared to nonsymbolic numerical magnitude judgments. The reverse comparison (nonsymbolic>symbolic) revealed several regions including the right posterior superior parietal lobe. These results reveal both format-general and format-specific processing of numerical stimuli in the brain. The potential roles of these regions in symbolic and nonsymbolic numerical processing are discussed.

Mapping numerical magnitudes onto symbols: the numerical distance effect and individual differences in children's mathematics achievement.

Although it is often assumed that abilities that reflect basic numerical understanding, such as numerical comparison, are related to children's mathematical abilities, this relationship has not been tested rigorously. In addition, the extent to which symbolic and nonsymbolic number processing play differential roles in this relationship is not yet understood. To address these questions, we collected mathematics achievement measures from 6- to 8-year-olds as well as reaction times from a numerical comparison task. Using the reaction times, we calculated the size of the numerical distance effect exhibited by each child. In a correlational analysis, we found that the individual differences in the distance effect were related to mathematics achievement but not to reading achievement. This relationship was found to be specific to symbolic numerical comparison. Implications for the role of basic numerical competency and the role of accessing numerical magnitude information from Arabic numerals for the development of mathematical skills and their impairment are discussed.

Domain-specific and domain-general changes in children's development of number comparison.

The numerical distance effect (inverse relationship between numerical distance and reaction time in relative number comparison tasks) has frequently been used to characterize the mental representation of number. The size of the distance effect decreases over developmental time. However, it is unclear whether this reduction simply reflects developmental changes in domain-general speed of processing and whether it is specific to numerical compared with non-numerical magnitude. To examine these open questions, we conducted a cross-sectional study with 6-, 7-, and 8-year-old children as well as adult college students. Participants performed comparisons on Arabic numerals, arrays of squares, squares of varying luminance and bars of varying height. To control for general age-related changes in reaction time, a measure of speed of processing was used as a covariate in the analysis. A significant developmental decrease in the distance effect was found across numerical and non-numerical comparison tasks over and above general changes in processing speed. However, this change was not found to differ as a function of format. These data suggest that developmental changes in the distance effect are reflective of changes in a domain-general comparison process, rather than domain-specific developmental changes in number representations. However, analysis of overall reaction times revealed significantly greater developmental changes for numerical relative to non-numerical comparison tasks. These findings highlight the importance of taking multiple measures into account when characterizing developmental changes in numerical magnitude processing. Implications for theories of numerical cognition and its development are discussed.

White matter microstructures underlying mathematical abilities in children.

The role of gray matter function and structure in mathematical cognition has been well researched. Comparatively little is known about white matter microstructures associated with mathematical abilities. Diffusion tensor imaging data from 13 children (7-9 years) and two measures of their mathematical competence were collected. Relationships between children's mathematical competence and fractional anisotropy were found in two left hemisphere white matter regions. Although the superior corona radiata was found to be associated with both numerical operations and mathematical reasoning, the inferior longitudinal fasciculus was correlated with numerical operations specifically. These findings suggest a role for microstructure in left white matter tracts for the development of mathematical skills. Moreover, the findings point to the involvement of different white matter tracts for numerical operations and mathematical reasoning.