Identifying conceptual neural responses to symbolic numerals.

The goal of measuring conceptual processing in numerical cognition is distanced by the possibility that neural responses to symbolic numerals are influenced by physical stimulus confounds. Here, we targeted conceptual responses to parity (even versus odd), using electroencephalogram (EEG) frequency-tagging with a symmetry/asymmetry design. Arabic numerals (2-9) were presented at 7.5 Hz in 50 s sequences; odd and even numbers were alternated to target differential, 'asymmetry' responses to parity at 3.75 Hz (7.5 Hz/2). Parity responses were probed with four different stimulus sets, increasing in intra-numeral stimulus variability, and with two control conditions composed of non-conceptual numeral alternations. Significant asymmetry responses were found over the occipitotemporal cortex to all conditions, even for the arbitrary controls. The large physical-differences control condition elicited the largest response in the stimulus set with the lowest variability (one font). Only in the stimulus set with the highest variability (20 drawn, coloured exemplars/numeral) did the response to parity surpass both control conditions. These findings show that physical differences across small sets of Arabic numerals can strongly influence, and even account for, automatic brain responses. However, carefully designed control conditions and highly variable stimulus sets may be used towards identifying truly conceptual neural responses.

Eye movements reveal that young school children shift attention when solving additions and subtractions.

Adults shift their attention to the right or to the left along a spatial continuum when solving additions and subtractions, respectively. Studies suggest that these shifts not only support the exact computation of the results but also anticipatively narrow down the range of plausible answers when processing the operands. However, little is known on when and how these attentional shifts arise in childhood during the acquisition of arithmetic. Here, an eye-tracker with high spatio-temporal resolution was used to measure spontaneous eye movements, used as a proxy for attentional shifts, while children of 2nd (8 y-o; N = 50) and 4th (10 y-o; N = 48) Grade solved simple additions (e.g., 4+3) and subtractions (e.g., 3-2). Gaze patterns revealed horizontal and vertical attentional shifts in both groups. Critically, horizontal eye movements were observed in 4th Graders as soon as the first operand and the operator were presented and thus before the beginning of the exact computation. In 2nd Graders, attentional shifts were only observed after the presentation of the second operand just before the response was made. This demonstrates that spatial attention is recruited when children solve arithmetic problems, even in the early stages of learning mathematics. The time course of these attentional shifts suggests that with practice in arithmetic children start to use spatial attention to anticipatively guide the search for the answer and facilitate the implementation of solving procedures. RESEARCH HIGHLIGHTS: Additions and subtractions are associated to right and left attentional shifts in adults, but it is unknown when these mechanisms arise in childhood. Children of 8-10 years old solved single-digit additions and subtractions while looking at a blank screen. Eye movements showed that children of 8 years old already show spatial biases possibly to represent the response when knowing both operands. Children of 10 years old shift attention before knowing the second operand to anticipatively guide the search for plausible answers.

The neural signature of numerosity by separating numerical and continuous magnitude extraction in visual cortex with frequency-tagged EEG.

The ability to handle approximate quantities, or number sense, has been recurrently linked to mathematical skills, although the nature of the mechanism allowing to extract numerical information (i.e., numerosity) from environmental stimuli is still debated. A set of objects is indeed not only characterized by its numerosity but also by other features, such as the summed area occupied by the elements, which often covary with numerosity. These intrinsic relations between numerosity and nonnumerical magnitudes led some authors to argue that numerosity is not independently processed but extracted through a weighting of continuous magnitudes. This view cannot be properly tested through classic behavioral and neuroimaging approaches due to these intrinsic correlations. The current study used a frequency-tagging EEG approach to separately measure responses to numerosity as well as to continuous magnitudes. We recorded occipital responses to numerosity, total area, and convex hull changes but not to density and dot size. We additionally applied a model predicting primary visual cortex responses to the set of stimuli. The model output was closely aligned with our electrophysiological data, since it predicted discrimination only for numerosity, total area, and convex hull. Our findings thus demonstrate that numerosity can be independently processed at an early stage in the visual cortex, even when completely isolated from other magnitude changes. The similar implicit discrimination for numerosity as for some continuous magnitudes, which correspond to basic visual percepts, shows that both can be extracted independently, hence substantiating the nature of numerosity as a primary feature of the visual scene.

Harmonic Amplitude Summation for Frequency-tagging Analysis.

In the approach of frequency tagging, stimuli that are presented periodically generate periodic responses of the brain. Following a transformation into the frequency domain, the brain's response is often evident at the frequency of stimulation, F, and its higher harmonics (2F, 3F, etc.). This approach is increasingly used in neuroscience, as it affords objective measures to characterize brain function. However, whether these specific harmonic frequency responses should be combined for analysis-and if so, how-remains an outstanding issue. In most studies, higher harmonic responses have not been described or were described only individually; in other studies, harmonics have been combined with various approaches, for example, averaging and root-mean-square summation. A rationale for these approaches in the context of frequency-based analysis principles and an understanding of how they relate to the brain's response amplitudes in the time domain have been missing. Here, with these elements addressed, the summation of (baseline-corrected) harmonic amplitude is recommended.

Developmental changes in neural letter-selectivity: A 1-year follow-up of beginning readers.

The developmental course of neural tuning to visual letter strings is unclear. Here we tested 39 children longitudinally, at the beginning of grade 1 (6.45 ± 0.33 years old) and 1 year after, with fast periodic visual stimulation in electroencephalography to assess the evolution of selective neural responses to letter strings and their relationship with emerging reading abilities. At both grades, frequency-tagged letter strings were discriminated from pseudofont strings (i.e. letter-selectivity) over the left occipito-temporal cortex, with effects observed at the individual level in 62% of children. However, visual words were not discriminated from pseudowords (lexical access) at either grade. Following 1 year of schooling, letter-selective responses showed a specific increase in amplitude, a more complex pattern of harmonics, and were located more anteriorly over the left occipito-temporal cortex. Remarkably, at both grades, neural responses were highly significant at the individual level and correlated with individual reading scores. The amplitude increase in letter-selective responses between grades was not found for discrimination responses of familiar keyboard symbols from pseudosymbols, and was not related to a general increase in visual stimulation responses. These findings demonstrate a rapid onset of left hemispheric letter selectivity, with 1 year of reading instruction resulting in increased emerging reading abilities and a clear quantitative and qualitative evolution within left hemispheric neural circuits for reading.

The importance of visuospatial abilities for verbal number skills in preschool: Adding spatial language to the equation.

Children's verbal number skills set the foundation for mathematical development. Therefore, it is central to understand their cognitive origins. Evidence suggests that preschool children rely on visuospatial abilities when solving counting and number naming tasks despite their predominantly verbal nature. We aimed to replicate these findings when controlling for verbal abilities and sociodemographic factors. Moreover, we further characterized the relation between visuospatial abilities and verbal number skills by examining the role of spatial language. Because spatial language encompasses the verbalization of spatial thinking, it is a key candidate supporting the interplay between visuospatial and verbal processes. Regression analysis indicated that both visuospatial and verbal abilities, as assessed by spatial perception and phonological awareness, respectively, uniquely predicted verbal number skills when controlling for their respective influences, age, gender, and socioeconomic status. This confirms the spatial grounding of verbal number skills. Interestingly, adding spatial language to the model abolished the predictive effects of visuospatial and verbal abilities, whose influences were completely mediated by spatial language. Verbal number skills thus concurrently depend on specifically those visuospatial and verbal processes jointly indexed through spatial language. The knowledge of spatial terms might promote verbal number skills by advancing the understanding of the spatial relations between numerical magnitudes on the mental number line. Promoting spatial language in preschool thus might be a successful avenue for stimulating mathematical development prior to formal schooling. Moreover, measures of spatial language could become an additional promising tool to screen preschool children for potential upcoming difficulties with mathematical learning.

The right hemispheric dominance for face perception in preschool children depends on the visual discrimination level.

The developmental origin of human adults' right hemispheric dominance in response to face stimuli remains unclear, in particular because young infants' right hemispheric advantage in face-selective response is no longer present in preschool children, before written language acquisition. Here we used fast periodic visual stimulation (FPVS) with scalp electroencephalography (EEG) to test 52 preschool children (5.5 years old) at two different levels of face discrimination: discrimination of faces against objects, measuring face-selectivity, or discrimination between individual faces. While the contrast between faces and nonface objects elicits strictly bilateral occipital responses in children, strengthening previous observations, discrimination of individual faces in the same children reveals a strong right hemispheric lateralization over the occipitotemporal cortex. Picture-plane inversion of the face stimuli significantly decreases the individual discrimination response, although to a much smaller extent than in older children and adults tested with the same paradigm. However, there is only a nonsignificant trend for a decrease in right hemispheric lateralization with inversion. There is no relationship between the right hemispheric lateralization in individual face discrimination and preschool levels of readings abilities. The observed difference in the right hemispheric lateralization obtained in the same population of children with two different paradigms measuring neural responses to faces indicates that the level of visual discrimination is a key factor to consider when making inferences about the development of hemispheric lateralization of face perception in the human brain.

Lateralized Neural Responses to Letters and Digits in First Graders.

The emergence of visual cortex specialization for culturally acquired characters like letters and digits, both arbitrary shapes related to specific cognitive domains, is yet unclear. Here, 20 young children (6.12 years old) were tested with a frequency-tagging paradigm coupled with electroencephalogram recordings to assess discrimination responses of letters from digits and vice-versa. One category of stimuli (e.g., letters) was periodically inserted (1/5) in streams of the other category (e.g., digits) presented at a fast rate (6 Hz). Results show clear right-lateralized discrimination responses at 6 Hz/5 for digits within letters, and a trend for left-lateralization for letters. These results support an early developmental emergence of ventral occipito-temporal cortex specialization for visual recognition of digits and letters, potentially in relation with relevant coactivated brain networks.

Visuo-spatial abilities are key for young children's verbal number skills.

Children's development of verbal number skills (i.e., counting abilities and knowledge of the number names) presents a milestone in mathematical development. Different factors such as visuo-spatial and verbal abilities have been discussed as contributing to the development of these foundational skills. To understand the cognitive nature of verbal number skills in young children, the current study assessed the relation of preschoolers' verbal and visuo-spatial abilities to their verbal number skills. In total, 141 children aged 5 or 6 years participated in the current study. Verbal number skills were regressed on vocabulary, phonological awareness and visuo-spatial abilities, and verbal and visuo-spatial working memory in a structural equation model. Only visuo-spatial abilities emerged as a significant predictor of verbal number skills in the estimated model. Our results suggest that visuo-spatial abilities contribute to a larger extent to children's verbal number skills than verbal abilities. From a theoretical point of view, these results suggest a visuo-spatial, rather than a verbal, grounding of verbal number skills. These results are potentially informative for the conception of early mathematics assessments and interventions.

Mathematical abilities in elementary school: Do they relate to number-space associations?

Considering the importance of mathematics in Western societies, it is crucial to understand the cognitive processes involved in the acquisition of more complex mathematical skills. The current study, therefore, investigated how the quality of number-space mappings on the mental number line, as indexed by the parity SNARC (spatial-numerical association of response codes) effect, relates to mathematical performances in third- and fourth-grade elementary school children. Mathematical competencies were determined using the "Heidelberger Rechentest," a standardized German math test assessing both arithmetical and visuospatial math components. Stronger parity SNARC effects significantly related to better arithmetical but not visuospatial math abilities, albeit only in the relatively younger children. These findings highlight the importance of spatial-numerical interactions for arithmetical (as opposed to visuospatial) math skills at the fairly early stages of mathematical development. Differential relations might be explained by the reliance on problem-solving strategies involving number-space mappings only for arithmetic tasks mainly in younger children.

Speaking two languages with different number naming systems: What implications for magnitude judgments in bilinguals at different stages of language acquisition?

Differences between languages in terms of number naming systems may lead to performance differences in number processing. The current study focused on differences concerning the order of decades and units in two-digit number words (i.e., unit-decade order in German but decade-unit order in French) and how they affect number magnitude judgments. Participants performed basic numerical tasks, namely two-digit number magnitude judgments, and we used the compatibility effect (Nuerk et al. in Cognition 82(1):B25-B33, 2001) as a hallmark of language influence on numbers. In the first part we aimed to understand the influence of language on compatibility effects in adults coming from German or French monolingual and German-French bilingual groups (Experiment 1). The second part examined how this language influence develops at different stages of language acquisition in individuals with increasing bilingual proficiency (Experiment 2). Language systematically influenced magnitude judgments such that: (a) The spoken language(s) modulated magnitude judgments presented as Arabic digits, and (b) bilinguals' progressive language mastery impacted magnitude judgments presented as number words. Taken together, the current results suggest that the order of decades and units in verbal numbers may qualitatively influence magnitude judgments in bilinguals and monolinguals, providing new insights into how number processing can be influenced by language(s).

Are patients with Parkinson's disease blind to blindsight?

In Parkinson's disease, visual dysfunction is prominent. Visual hallucinations can be a major hallmark of late stage disease, but numerous visual deficits also occur in early stage Parkinson's disease. Specific retinopathy, deficits in the primary visual pathway and the secondary ventral and dorsal pathways, as well as dysfunction of the attention pathways have all been posited as causes of hallucinations in Parkinson's disease. We present data from patients with Parkinson's disease that contrast with a known neuro-ophthalmological syndrome, termed 'blindsight'. In this syndrome, there is an absence of conscious object identification, but preserved 'guess' of the location of a stimulus, preserved reflexive saccades and motion perception and preserved autonomical and expressive reactions to negative emotional facial expressions. We propose that patients with Parkinson's disease have the converse of blindsight, being 'blind to blindsight'. As such they preserve conscious vision, but show erroneous 'guess' localization of visual stimuli, poor saccades and motion perception, and poor emotional face perception with blunted autonomic reaction. Although a large data set on these deficits in Parkinson's disease has been accumulated, consolidation into one specific syndrome has not been proposed. Focusing on neuropathological and physiological data from two phylogenetically old and subconscious pathways, the retino-colliculo-thalamo-amygdala and the retino-geniculo-extrastriate pathways, we propose that aberrant function of these systems, including pathologically inhibited superior colliculus activity, deficient corollary discharges to the frontal eye fields, dysfunctional pulvinar, claustrum and amygdaloid subnuclei of the amygdala, the latter progressively burdened with Lewy bodies, underlie this syndrome. These network impairments are further corroborated by the concept of the 'silent amygdala'. Functionally being 'blind to blindsight' may facilitate the highly distinctive 'presence' or 'passage' hallucinations of Parkinson's disease and can help to explain handicaps in driving capacities and dysfunctional 'theory of mind'. We propose this synthesis to prompt refined neuropathological and neuroimaging studies on the pivotal nuclei in these pathways in order to better understand the networks underpinning this newly conceptualized syndrome in Parkinson's disease.

Face perception is tuned to horizontal orientation in the N170 time window.

The specificity of face perception is thought to reside both in its dramatic vulnerability to picture-plane inversion and its strong reliance on horizontally oriented image content. Here we asked when in the visual processing stream face-specific perception is tuned to horizontal information. We measured the behavioral performance and scalp event-related potentials (ERP) when participants viewed upright and inverted images of faces and cars (and natural scenes) that were phase-randomized in a narrow orientation band centered either on vertical or horizontal orientation. For faces, the magnitude of the inversion effect (IE) on behavioral discrimination performance was significantly reduced for horizontally randomized compared to vertically or nonrandomized images, confirming the importance of horizontal information for the recruitment of face-specific processing. Inversion affected the processing of nonrandomized and vertically randomized faces early, in the N170 time window. In contrast, the magnitude of the N170 IE was much smaller for horizontally randomized faces. The present research indicates that the early face-specific neural representations are preferentially tuned to horizontal information and offers new perspectives for a description of the visual information feeding face-specific perception.

The impact of inhibition capacities and age on number-space associations.

Numerical and spatial representations are tightly linked, i.e., when doing a binary classification judgment on Arabic digits, participants are faster to respond with their left/right hand to small/large numbers, respectively (Spatial-Numerical Association of Response Codes, SNARC effect, Dehaene et al. in J Exp Psychol Gen 122:371-396, 1993). To understand the underlying mechanisms of the well-established SNARC effect, it seems essential to explore the considerable inter-individual variability characterizing it. The present study assesses the respective roles of inhibition, age, working memory (WM) and response speed. Whereas these non-numerical factors have been proposed as potentially important factors to explain individual differences in SNARC effects, none (except response speed) has so far been explored directly. Confirming our hypotheses, the results show that the SNARC effect was stronger in participants that had weaker inhibition abilities (as assessed by the Stroop task), were relatively older and had longer response times. Interestingly, whereas a significant part of the age influence was mediated by cognitive inhibition, age also directly impacted the SNARC effect. Similarly, cognitive inhibition abilities explained inter-individual variability in number-space associations over and above the factors age, WM capacity and response speed. Taken together our results provide new insights into the nature of number-space associations by describing how these are influenced by the non-numerical factors age and inhibition.

Weaker semantic priming effects with number words in the second language of math learning.

Bilinguals' exact number representations result from associations between language-independent Indo-Arabic digits ("5"), two verbal codes ("fünf" and "cinq") and a common, largely overlapping semantic representation. To compare the lexical and semantic access to number representations between two languages, we recruited a sample of balanced highly proficient German-French adult bilinguals. At school, those bilinguals learned mathematics in German for 6 years (LM1) and then switched to French (LM2) in 7th grade (12 years old) until 13th grade. After the brief presentation of primes (51 ms) consisting of Indo-Arabic digits or number words in German or French, an Indo-Arabic digits target had to be read in either German or French in an online study. Stimuli were numbers from 1 to 9, and we varied the absolute distance between primes and targets from 0 (i.e., 1-1) to 3 (1-4; as in Reynvoet et al., 2002). The priming distance effect (PDE) was used to measure the strength of numerical semantic association. We find comparable PDEs with Indo-Arabic digits and German number word primes, independently from the target naming language. However, we did not find a clear PDE with French number word primes, neither when naming targets in German, nor in French. The weaker PDE from LM2 compared to LM1 primes is interpreted as a weaker lexico-semantic association of LM2 number words. These results indicate a critical role of the LM1 and further emphasize the role of language in processing numbers. They might have important implications for designing bilingual school curricula. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Linguistic and attentional factors - Not statistical regularities - Contribute to word-selective neural responses with FPVS-oddball paradigms.

Studies using frequency-tagging in electroencephalography (EEG) have dramatically increased in the past 10 years, in a variety of domains and populations. Here we used Fast Periodic Visual Stimulation (FPVS) combined with an oddball design to explore visual word recognition. Given the paradigm's high sensitivity, it is crucial for future basic research and clinical application to prove its robustness across variations of designs, stimulus types and tasks. This paradigm uses periodicity of brain responses to measure discrimination between two experimentally defined categories of stimuli presented periodically. EEG was recorded in 22 adults who viewed words inserted every 5 stimuli (at 2 Hz) within base stimuli presented at 10 Hz. Using two discrimination levels (deviant words among nonwords or pseudowords), we assessed the impact of relative frequency of item repetition (set size or item repetition controlled for deviant versus base stimuli), and of the orthogonal task (focused or deployed spatial attention). Word-selective occipito-temporal responses were robust at the individual level (significant in 95% of participants), left-lateralized, larger for the prelexical (nonwords) than lexical (pseudowords) contrast, and stronger with a deployed spatial attention task as compared to the typically used focused task. Importantly, amplitudes were not affected by item repetition. These results help understanding the factors influencing word-selective EEG responses and support the validity of FPVS-EEG oddball paradigms, as they confirm that word-selective responses are linguistic. Second, they show its robustness against design-related factors that could induce statistical (ir)regularities in item rate. They also confirm its high individual sensitivity and demonstrate how it can be optimized, using a deployed rather than focused attention task, to measure implicit word recognition processes in typical and atypical populations.

Developing number-space associations: SNARC effects using a color discrimination task in 5-year-olds.

Human adults' numerical representation is spatially oriented; consequently, participants are faster to respond to small/large numerals with their left/right hand, respectively, when doing a binary classification judgment on numbers, known as the SNARC (spatial-numerical association of response codes) effect. Studies on the emergence and development of the SNARC effect remain scarce. The current study introduces an innovative new paradigm based on a simple color judgment of Arabic digits. Using this task, we found a SNARC effect in children as young as 5.5 years. In contrast, when preschool children needed to perform a magnitude judgment task necessitating exact number knowledge, the SNARC effect started to emerge only at 5.8 years. Moreover, the emergence of a magnitude SNARC but not a color SNARC was linked to proficiency with Arabic digits. Our results suggest that access to a spatially oriented approximate magnitude representation from symbolic digits emerges early in ontogenetic development. Exact magnitude judgments, on the other hand, rely on experience with Arabic digits and, thus, necessitate formal or informal schooling to give access to a spatially oriented numerical representation.

The importance of spatial language for early numerical development in preschool: Going beyond verbal number skills.

Recent evidence suggests that spatial language in preschool positively affects the development of verbal number skills, as indexed by aggregated performances on counting and number naming tasks. We firstly aimed to specify whether spatial language (the knowledge of locative prepositions) significantly relates to both of these measures. In addition, we assessed whether the predictive value of spatial language extends beyond verbal number skills to numerical subdomains without explicit verbal component, such as number writing, symbolic magnitude classifications, ordinal judgments and numerosity comparisons. To determine the unique contributions of spatial language to these numerical skills, we controlled in our regression analyses for intrinsic and extrinsic spatial abilities, phonological awareness as well as age, socioeconomic status and home language. With respect to verbal number skills, it appeared that spatial language uniquely predicted forward and backward counting but not number naming, which was significantly affected only by phonological awareness. Regarding numerical tasks that do not contain explicit verbal components, spatial language did not relate to number writing or numerosity comparisons. Conversely, it explained unique variance in symbolic magnitude classifications and was the only predictor of ordinal judgments. These findings thus highlight the importance of spatial language for early numerical development beyond verbal number skills and suggest that the knowledge of spatial terms is especially relevant for processing cardinal and ordinal relations between symbolic numbers. Promoting spatial language in preschool might thus be an interesting avenue for fostering the acquisition of these symbolic numerical skills prior to formal schooling.

Categorical consistency facilitates implicit learning of color-number associations.

In making sense of the environment, we implicitly learn to associate stimulus attributes that frequently occur together. Is such learning favored for categories over individual items? Here, we introduce a novel paradigm for directly comparing category- to item-level learning. In a category-level experiment, even numbers (2,4,6,8) had a high-probability of appearing in blue, and odd numbers (3,5,7,9) in yellow. Associative learning was measured by the relative performance on trials with low-probability (p = .09) to high-probability (p = .91) number colors. There was strong evidence for associative learning: low-probability performance was impaired (40ms RT increase and 8.3% accuracy decrease relative to high-probability). This was not the case in an item-level experiment with a different group of participants, in which high-probability colors were non-categorically assigned (blue: 2,3,6,7; yellow: 4,5,8,9; 9ms RT increase and 1.5% accuracy increase). The categorical advantage was upheld in an explicit color association report (83% accuracy vs. 43% at the item-level). These results support a conceptual view of perception and suggest empirical bases of categorical, not item-level, color labeling of learning materials.