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2009, Cortex
Healthy human brains come equipped with several circuits that contribute to number processing. Nature and nurture interact to produce a unique combination of core skills and more sophisticated abilities, by building on a handful of auxiliary routes (e.g., verbal language, body knowledge and visuospatial attention). Transcranial magnetic stimulation (TMS) studies on number processing will be here succinctly reviewed, in light of their most stimulating and challenging contributions. New research directions will be pointed out, that might enhance their theoretical impact. (M. Sandrini). a v a i l a b l e a t w w w . s c i e n c e d i r e c t . c o m j o u r n a l h o m e p a g e : w w w . e l s e v i e r . c o m / l o c a t e / c o r t e x ARTICLE IN PRESS 0010-9452/$ -see front matter ª
Frontiers in Human Neuroscience, 2011
How numerical representation is encoded in the adult human brain is important for a basic understanding of human brain organization, its typical and atypical development, its evolutionary precursors, cognitive architectures, education, and rehabilitation. Previous studies have shown that numerical processing activates the same intraparietal regions irrespective of the presentation format (e.g., symbolic digits or non-symbolic dot arrays). This has led to claims that there is a single format-independent, numerical representation. In the current study we used a functional magnetic resonance adaptation paradigm, and effective connectivity analysis to re-examine whether numerical processing in the intraparietal sulci is dependent or independent on the format of the stimuli. We obtained two novel results. First, the whole brain analysis revealed that format change (e.g., from dots to digits), in the absence of a change in magnitude, activated the same intraparietal regions as magnitude change, but to a greater degree. Second, using dynamic causal modeling as a tool to disentangle neuronal specialization across regions that are commonly activated, we found that the connectivity between the left and right intraparietal sulci is format-dependent. Together, this line of results supports the idea that numerical representation is subserved by multiple mechanisms within the same parietal regions.
Abstract In adult human brains, the horizontal segment of the intraparietal sulcus plays a large role in representing numeric magnitude. In children and non-human primates, however, frontal cortex may play a larger role. We hypothesized that there is a link between observed developmental changes in locus of representation (frontal to parietal) and type of representation used (logarithmic to linear). Participants were presented with number lines and asked to judge accuracy of linear, logarithmic, or log-linear placements.
Journal of Clinical Medicine
Atypical development of numerical cognition (dyscalculia) may increase the onset of neuropsychiatric symptoms, especially when untreated, and it may have long-term detrimental social consequences. However, evidence-based treatments are still lacking. Despite plenty of studies investigating the effects of transcranial electrical stimulation (tES) on numerical cognition, a systematized synthesis of results is still lacking. In the present systematic review (PROSPERO ID: CRD42021271139), we found that the majority of reports (20 out of 26) showed the effectiveness of tES in improving both number (80%) and arithmetic (76%) processing. In particular, anodal tDCS (regardless of lateralization) over parietal regions, bilateral tDCS (regardless of polarity/lateralization) over frontal regions, and tRNS (regardless of brain regions) strongly enhance number processing. While bilateral tDCS and tRNS over parietal and frontal regions and left anodal tDCS over frontal regions consistently improv...
Journal of Cognitive …, 2007
Developmental and cross-cultural studies show that finger-counting represents one of the basic number learning strategies. However, despite the ubiquity of such an embodied strategy, the issue of whether there is a neural link between numbers and fingers in adult, literate individuals remains debated. Here, we used transcranial magnetic stimulation to study changes of excitability of hand muscles of individuals performing a visual parity judgment task, a task not requiring counting, on Arabic numerals from 1 to 9. While no modulation was observed for the left hand muscles, an increase in amplitude of motor evoked potentials was found for the right hand muscles. This increase was specific for smaller numbers (1 to 4) as compared to larger numbers (6 to 9). These findings indicate a close relationship between hand/finger and numerical representations.
Frontiers in human neuroscience, 2013
A central question in cognitive and educational neuroscience is whether brain operations supporting nonlinguistic intuitive number sense (numerosity) predict individual acquisition and academic achievement for symbolic or "formal" math knowledge. Here, we conducted a developmental functional magnetic resonance imaging (MRI) study of nonsymbolic numerosity task performance in 44 participants including 14 school age children (6-12 years old), 14 adolescents (13-17 years old), and 16 adults and compared a brain activity measure of numerosity precision to scores from the Woodcock-Johnson III Broad Math index of math academic achievement. Accuracy and reaction time from the numerosity task did not reliably predict formal math achievement. We found a significant positive developmental trend for improved numerosity precision in the parietal cortex and intraparietal sulcus specifically. Controlling for age and overall cognitive ability, we found a reliable positive relationship between individual math achievement scores and parietal lobe activity only in children. In addition, children showed robust positive relationships between math achievement and numerosity precision within ventral stream processing areas bilaterally. The pattern of results suggests a dynamic developmental trajectory for visual discrimination strategies that predict the acquisition of formal math knowledge. In adults, the efficiency of visual discrimination marked by numerosity acuity in ventral occipital-temporal cortex and hippocampus differentiated individuals with better or worse formal math achievement, respectively. Overall, these results suggest that two different brain systems for nonsymbolic numerosity acuity may contribute to individual differences in math achievement and that the contribution of these systems differs across development. Hum Brain Mapp 36:804-826, 2015. V C 2014 Wiley Periodicals, Inc. in Wiley Online Library (wileyonlinelibrary.com). r Human Brain Mapping 36:804-826 (2015) r V C 2014 Wiley Periodicals, Inc. r Numerosity Development and Math Achievement r r 805 r r Haist et al r r 806 r r Numerosity Development and Math Achievement r r 807 r r Haist et al r r 808 r r Haist et al r r 810 r r Haist et al r r 824 r
Neuropsychologia, 2017
It is currently debated whether numbers are processed using a number-specific system or a general magnitude processing system, also used for non-numerical magnitudes such as physical size, duration, or luminance. Activation likelihood estimation (ALE) was used to conduct the first quantitative meta-analysis of 93 empirical neuroimaging papers examining neural activation during numerical and non-numerical magnitude processing. Foci were compiled to generate probabilistic maps of activation for non-numerical magnitudes (e.g. physical size), symbolic numerical magnitudes (e.g. Arabic digits), and nonsymbolic numerical magnitudes (e.g. dot arrays). Conjunction analyses revealed overlapping activation for symbolic, nonsymbolic and non-numerical magnitudes in frontal and parietal lobes. Contrast analyses revealed specific activation in the left superior parietal lobule for symbolic numerical magnitudes. In contrast, small regions in the bilateral precuneus were specifically activated for ...
Journal of Cognitive Neuroscience, 2014
A dominant hypothesis on how the brain processes numerical size proposes a spatial representation of numbers as positions on a 'mental number line'. An alternative hypothesis considers numbers as elements of a generalized representation of sensorimotor-related magnitude which is not obligatorily spatial. Here we show that individuals' relative use of spatial and non-spatial representations has a cerebral counterpart in the structural organization of the posterior parietal cortex. Inter-individual variability in the linkage between numbers and spatial responses (faster left responses to low numbers and right responses to high numbers; SNARC effect) correlated with variations in grey matter volume around the right precuneus. Conversely, differences in the disposition to link numbers to force production (faster soft responses to low numbers and hard responses to high numbers) were related to grey matter volume in the left angular gyrus. This finding suggests that numerical cognition relies on multiple mental representations of analogue magnitude using different neural implementations that are linked to individual traits.
Neuroscience Letters, 2009
Current Biology, 2004
The parietal cortex is a central part of the brain's system for representing numbers and magnitudes. Activity in the parietal cortex might reflect number representation or actions made in response to the numbers.
Journal of Cognitive Neuroscience, 2008
Annual Review of Neuroscience, 2009
Number symbols have allowed humans to develop superior mathematical skills that are a hallmark of technologically advanced cultures. Findings in animal cognition, developmental psychology, and anthropology indicate that these numerical skills are rooted in nonlinguistic biological primitives. Recent studies in human and nonhuman primates using a broad range of methodologies provide evidence that numerical information is represented and processed by regions of the prefrontal and posterior parietal lobes, with the intraparietal sulcus as a key node for the representation of the semantic aspect of numerical quantity.
Nature Reviews Neuroscience, 2008
NeuroImage, 2001
To investigate the hemispheric organization of a language-independent spatial representation of number magnitude in the human brain we applied focal repetitive transcranial magnetic stimulation (rTMS) to the right or left angular gyrus while subjects performed a number comparison task with numbers between 31 and 99. Repetitive TMS over the angular gyrus disrupted performance of a visuospatial search task, and rTMS at the same site disrupted organization of the putative "number line." In some cases the pattern of disruption caused by angular gyrus rTMS suggested that this area normally mediates a spatial representation of number. The effect of angular gyrus rTMS on the number line task was specific. rTMS had no disruptive effect when delivered over another parietal region, the supramarginal gyrus, in either the left or the right hemisphere.
2020
Humans and other animal species are endowed with the ability to sense, represent, and mentally manipulate the number of items in a set without needing to count them. One central hypothesis is that this ability relies on an automated functional system dedicated to numerosity, the perception of the discrete numerical magnitude of a set of items. This system has classically been associated with intraparietal regions, however accumulating evidence in favor of an early visual number sense calls into question the functional role of parietal regions in numerosity processing. Targeting specifically numerosity among other visual features in the earliest stages of processing requires high temporal and spatial resolution. We used frequency-tagged magnetoencephalography (MEG) to investigate the early automatic processing of numerical magnitudes and measured the steady-state brain responses specifically evoked by numerical and other visual changes in the visual scene. The neuromagnetic responses...
Cerebral Cortex, 2010
Numerous studies have identified the intraparietal sulcus (IPS) as an area critically involved in numerical processing. IPS neurons in macaques are tuned to a preferred numerosity, hence neurally coding numerosity in a number-selective way. Neuroimaging studies in humans have demonstrated number-selective processing in the anterior parts of the IPS. Nevertheless, the processes that convert visual input into a number-selective neural code remain unknown. Computational studies have suggested that a neural coding stage that is sensitive, but not selective to number, precedes numberselective coding when processing nonsymbolic quantities but not when processing symbolic quantities. In Experiment 1, we used functional magnetic resonance imaging to localize number-sensitive areas in the human brain by searching for areas exhibiting increasing activation with increasing number, carefully controlling for nonnumerical parameters. An area in posterior superior parietal cortex was identified as a substrate for the intermediate number-sensitive steps required for processing nonsymbolic quantities. In Experiment 2, the interpretation of Experiment 1 was confirmed with a connectivity analysis showing that a shared number-selective representation in IPS is reached through different pathways for symbolic versus nonsymbolic quantities. The preferred pathway for processing nonsymbolic quantities included the number-sensitive area in superior parietal cortex, whereas the pathway for processing symbolic quantities did not.
PsycEXTRA Dataset
A dominant hypothesis on how the brain processes numerical size proposes a spatial representation of numbers as positions on a 'mental number line'. An alternative hypothesis considers numbers as elements of a generalized representation of sensorimotor-related magnitude which is not obligatorily spatial. Here we show that individuals' relative use of spatial and non-spatial representations has a cerebral counterpart in the structural organization of the posterior parietal cortex. Inter-individual variability in the linkage between numbers and spatial responses (faster left responses to low numbers and right responses to high numbers; SNARC effect) correlated with variations in grey matter volume around the right precuneus. Conversely, differences in the disposition to link numbers to force production (faster soft responses to low numbers and hard responses to high numbers) were related to grey matter volume in the left angular gyrus. This finding suggests that numerical cognition relies on multiple mental representations of analogue magnitude using different neural implementations that are linked to individual traits.
Trends in Neurosciences, 1998
). The number domain is a prime example where strong evidence points to an evolutionary endowment of abstract domain-specific knowledge in the brain because there are parallels between number processing in animals and humans.The numerical distance effect, which refers to the finding that the ability to discriminate between two numbers improves as the numerical distance between them increases, has been demonstrated in humans and animals, as has the number size effect, which refers to the finding that for equal numerical distance,discrimination of two numbers worsens as their numerical size increases.
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