The neural circuits of number and letter copying: an fNIRS study

In our daily lives, we are constantly exposed to numbers and letters. However, it is still under debate how letters and numbers are processed in the brain, while information on this topic would allow for a more comprehensive understanding of, for example, known influences of language on numerical cognition or neural circuits shared by numerical cognition and language processing. Some findings provide evidence for a double dissociation between numbers and letters, with numbers being represented in the right and letters in the left hemisphere, while the opposing view suggests a shared neural network. Since processing may depend on the task, we address the reported inconsistencies in a very basic symbol copying task using functional near-infrared spectroscopy (fNIRS). fNIRS data revealed that both number and letter copying rely on the bilateral middle and left inferior frontal gyri. Only numbers elicited additional activation in the bilateral parietal cortex and in the left superior temporal gyrus. However, no cortical activation difference was observed between copying numbers and letters, and there was Bayesian evidence for common activation in the middle frontal gyri and superior parietal lobules. Therefore, we conclude that basic number and letter processing are based on a largely shared cortical network, at least in a simple task such as copying symbols. This suggests that copying can be used as a control condition for more complex tasks in neuroimaging studies without subtracting stimuli-specific activation

A Neural Model of How The Brain Represents and Compares Numbers

Many psychophysical experiments have shown that the representation of numbers and numerical quantities in humans and animals is related to number magnitude. A neural network model is proposed to quantitatively simulate error rates in quantification and numerical comparison tasks, and reaction times for number priming and numerical assessment and comparison tasks. Transient responses to inputs arc integrated before they activate an ordered spatial map that selectively responds to the number of events in a sequence. The dynamics of numerical comparison are encoded in activity pattern changes within this spatial map. Such changes cause a "directional comparison wave" whose properties mimic data about numerical comparison. These model mechanisms are variants of neural mechanisms that have elsewhere been used to explain data about motion perception, attention shifts, and target tracking. Thus, the present model suggests how numerical representations may have emerged as specializations of more primitive mechanisms in the cortical Where processing stream.National Science Foundation (IRI-97-20333); Defense Advanced research Projects Agency and the Office of Naval Research (N00014-95-1-0409); National Institute of Health (1-R29-DC02952-01

Early dissociation of numbers and letters in the human brain

Published online 7 May 2020Numbers and letters are culturally created symbols which are learned through repeated training. This experience leads to a functional specialization of the perceptual system of our brain. Recent evidence suggests a neural dissociation between these two symbols. While previous literature has shown that letters elicit a left lateralized neural response, new studies suggest that numbers elicit preferentially a bilateral or right lateralized response. However, the time course of the neural patterns that characterize this dissociation is still underspecified. In the present study, we investigated with magnetoencephalography (MEG) the spatio-temporal dynamics of the neural response generated by numbers, letters and perceptually matched false fonts presented visually. Twenty-five healthy adults were recorded while participants performed a dot detection task. By including two experiments, we were able to study the effects of single characters as well as those of strings of characters. The signal analysis was focused on the event related fields (ERF) of the MEG signal in the sensors and in the source space. The main results of our study showed an early (<200 msec) preferential dissociation between single numbers and single letters on occipito-temporal sensors. When comparing strings of numbers and pseudowords, they differed also over prefrontal regions of the brain. These data offer a new example of acquired category-specific responses in the human brain.The research was partially supported by Basque Government (BERC 2018-2021 program), BCBL Severo Ochoa excellence accreditation SEV-2015-0490, and Grant RTI2018-093547-B-I00 from the Spanish Ministerio de Ciencia, Innovaci on y Universidades and the Agencia Estatal de Investigaci on

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

A Neural Model of How the Brain Represents and Compares Multi-Digit Numbers: Spatial and Categorical Processes

Both animals and humans are capable of representing and comparing numerical quantities, but only humans seem to have evolved multi-digit place-value number systems. This article develops a neural model, called the Spatial Number Network, or SpaN model, which predicts how these shared numerical capabilities are computed using a spatial representation of number quantities in the Where cortical processing stream, notably the Inferior Parietal Cortex. Multi-digit numerical representations that obey a place-value principle are proposed to arise through learned interactions between categorical language representations in the What cortical processing stream and the Where spatial representation. It is proposed that learned semantic categories that symbolize separate digits, as well as place markers like "tens," "hundreds," "thousands," etc., are associated through learning with the corresponding spatial locations of the Where representation, leading to a place-value number system as an emergent property of What-Where information fusion. The model quantitatively simulates error rates in quantification and numerical comparison tasks, and reaction times for number priming and numerical assessment and comparison tasks. In the Where cortical process, it is proposed that transient responses to inputs are integrated before they activate an ordered spatial map that selectively responds to the number of events in a sequence. Neural mechanisms are defined which give rise to an ordered spatial numerical map ordering and Weber law characteristics as emergent properties. The dynamics of numerical comparison are encoded in activity pattern changes within this spatial map. Such changes cause a "directional comparison wave" whose properties mimic data about numerical comparison. These model mechanisms are variants of neural mechanisms that have elsewhere been used to explain data about motion perception, attention shifts, and target tracking. Thus, the present model suggests how numerical representations may have emerged as specializations of more primitive mechanisms in the cortical Where processing stream. The model's What-Where interactions can explain human psychophysical data, such as error rates and reaction times, about multi-digit (base 10) numerical stimuli, and describe how such a competence can develop through learning. The SpaN model and its explanatory range arc compared with other models of numerical representation.Defense Advanced Research Projects Agency and the Office of Naval Research (N00014-95-1-0409); National Science Foundation (IRI-97-20333

Language and thought are not the same thing: evidence from neuroimaging and neurological patients

Is thought possible without language? Individuals with global aphasia, who have almost no ability to understand or produce language, provide a powerful opportunity to find out. Surprisingly, despite their near-total loss of language, these individuals are nonetheless able to add and subtract, solve logic problems, think about another person's thoughts, appreciate music, and successfully navigate their environments. Further, neuroimaging studies show that healthy adults strongly engage the brain's language areas when they understand a sentence, but not when they perform other nonlinguistic tasks such as arithmetic, storing information in working memory, inhibiting prepotent responses, or listening to music. Together, these two complementary lines of evidence provide a clear answer: many aspects of thought engage distinct brain regions from, and do not depend on, language

Experiential Effects on the Neural Substrates of Visual Word and Number Processing.

Visual processing of words and numbers is a uniquely human cognitive ability. Evidence suggests that a region in left occipitotemporal cortex, the so-called visual word form area (VWFA), is crucially involved in this ability, particularly in the visual recognition of words. In this dissertation, I present a methodological study and two empirical studies to investigate the role of experience in shaping the VWFA, to explore ways to estimate the amount of this experiential influence, and to examine how the neural substrates of visual number recognition are different from those of visual letter recognition. In the first study, I develop a novel statistical method to efficiently estimate correlation between paired spatial processes, and hence heritability in patterns of activation in neuroimaging datasets. The results demonstrate that the proposed method provides a better estimate of correlation and heritability than conventional voxelwise or region of interest methods. The second study applies this method in a monozygotic twin sample to explore the role of unique environmental effects in shaping VWFA activation. The results demonstrate that there are greater unique environmental effects for neural activity associated with familiar word recognition than with unfamiliar word recognition. The last study investigates whether the VWFA is also the crucial site for visual number recognition or whether number recognition is neurally dissociable from word recognition. I demonstrate letter-selective activation in left occipitotemporal cortex and number-selective activation in right lateral occipital cortex, thus establishing double dissociation. Furthermore, I show that individual differences in the laterality of visual number activation can be explained by individual differences in the laterality of numerical processing activation in parietal cortex. In sum, this dissertation investigates experiential effects on the neural substrates of visual word and number processing. In a methodological study, I present a more powerful statistical method to estimate correlation and heritability in neuroimaging datasets. The findings from the two empirical studies suggest a critical role of environment in shaping the VWFA, demonstrate a novel double dissociation between the neural substrates of letter and number recognition, and provide evidence that top-down influences give rise to the functional neural organization for visual number recognition.Ph.D.PsychologyUniversity of Michigan, Horace H. Rackham School of Graduate Studieshttp://deepblue.lib.umich.edu/bitstream/2027.42/86434/1/joonkoo_1.pd

Visual Occipito-Temporal N1 Sensitivity to Digits Across Elementary School

Number processing abilities are important for academic and personal development. The course of initial specialization of ventral occipito-temporal cortex (vOTC) sensitivity to visual number processing is crucial for the acquisition of numeric and arithmetic skills. We examined the visual N1, the electrophysiological correlate of vOTC activation across five time points in kindergarten (T1, mean age 6.60 years), middle and end of first grade (T2, 7.38 years; T3, 7.68 years), second grade (T4, 8.28 years), and fifth grade (T5, 11.40 years). A combination of cross-sectional and longitudinal EEG data of a total of 62 children (35 female) at varying familial risk for dyslexia were available to form groups of 23, 22, 27, 27, and 42 participants for each of the five time points. The children performed a target detection task which included visual presentation of single digits (DIG), false fonts (FF), and letters (LET) to derive measures for coarse (DIG vs. FF) and fine (DIG vs. LET) digit sensitive processing across development. The N1 amplitude analyses indicated coarse and fine sensitivity characterized by a stronger N1 to digits than false fonts across all five time points, and stronger N1 to digits than letters at all but the second (T2) time point. In addition, lower arithmetic skills were associated with stronger coarse N1 digit sensitivity over the left hemisphere in second grade (T4), possibly reflecting allocation of more attentional resources or stronger reliance on the verbal system in children with poorer arithmetic skills. To summarize, our results show persistent visual N1 sensitivity to digits that is already present early on in pre-school and remains stable until fifth grade. This pattern of digit sensitivity development clearly differs from the relatively sharp rise and fall of the visual N1 sensitivity to words or letters between kindergarten and middle of elementary school and suggests unique developmental trajectories for visual processing of written characters that are relevant to numeracy and literacy

Electrical cortical stimulation can impair production of the alphabet without impairing counting

Neurosurgery has the potential to cure patients with drug-resistant focal epilepsy, but carries the risk of permanent language impairment when surgery involves the dominant hemisphere of the brain. This risk can be estimated and minimized using electrical stimulation mapping (ESM), which uses cognitive and linguistic tasks during cortical ESM to differentiate "eloquent" and "resectable" areas in the brain. One such task, counting, is often used to screen and characterize language during ESM in patients whose language abilities are limited. Here we report a patient with drug-resistant epilepsy arising from the language-dominant hemisphere using fMRI. Our patient experienced loss of the ability to recite or write the alphabet, but not to count, during ESM of the dominant left posterior superior temporal gyrus. This selective impairment extended to both spoken and written production. We suggest the need for caution when using counting as a sole means to screen language function and as a method of testing low functioning patients using ESM