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.

Frequency-tagging EEG reveals the effect of attentional focus on abstract magnitude processing.

While humans can readily access the common magnitude of various codes such as digits, number words, or dot sets, it remains unclear whether this process occurs automatically, or only when explicitly attending to magnitude information. We addressed this question by examining the neural distance effect, a robust marker of magnitude processing, with a frequency-tagging approach. Electrophysiological responses were recorded while participants viewed rapid sequences of a base numerosity presented at 6 Hz (e.g., "2") in randomly mixed codes: digits, number words, canonical dot, and finger configurations. A deviant numerosity either close (e.g., "3") or distant (e.g., "8") from the base was inserted every five items. Participants were instructed to focus their attention either on the magnitude number feature (from a previous study), the parity number feature, a nonnumerical color feature or no specific feature. In the four attentional conditions, we found clear discrimination responses of the deviant numerosity despite its code variation. Critically, the distance effect (larger responses when base/deviant are distant than close) was present when participants were explicitly attending to magnitude and parity, but it faded with color and simple viewing instructions. Taken together, these results suggest automatic access to an abstract number representation but highlight the role of selective attention in processing the underlying magnitude information. This study therefore provides insights into how attention can modulate the neural activity supporting abstract magnitude processing.

Stronger neural response to canonical finger-number configurations in deaf compared to hearing adults revealed by FPVS-EEG.

The linguistic counting system of deaf signers consists of a manual counting format that uses specific structures for number words. Interestingly, the number signs from 1 to 4 in the Belgian sign languages correspond to the finger-montring habits of hearing individuals. These hand configurations could therefore be considered as signs (i.e., part of a language system) for deaf, while they would simply be number gestures (not linguistic) for hearing controls. A Fast Periodic Visual Stimulation design was used with electroencephalography recordings to examine whether these finger-number configurations are differently processed by the brain when they are signs (in deaf signers) as compared to when they are gestures (in hearing controls). Results showed that deaf signers show stronger discrimination responses to canonical finger-montring configurations compared to hearing controls. A second control experiment furthermore demonstrated that this finding was not merely due to the experience deaf signers have with the processing of hand configurations, as brain responses did not differ between groups for finger-counting configurations. Number configurations are therefore processed differently by deaf signers, but only when these configurations are part of their language system.

Spatial Resolution Evaluation Based on Experienced Visual Categories With Sweep Evoked Periodic EEG Activity.

Purpose: Visual function is typically evaluated in clinical settings with visual acuity (VA), a test requiring to behaviorally match or name optotypes such as tumbling E or Snellen letters. The ability to recognize these symbols has little in common with the automatic and rapid visual recognition of socially important stimuli in real life. Here we use sweep visual evoked potentials to assess spatial resolution objectively based on the recognition of human faces and written words. Methods: To this end, we tested unfamiliar face individuation1 and visual word recognition2 in 15 normally sighted adult volunteers with a 68-electrode electroencephalogram system. Results: Unlike previous measures of low-level visual function including VA, the most sensitive electrode was found at an electrode different from Oz in a majority of participants. Thresholds until which faces and words could be recognized were evaluated at the most sensitive electrode defined individually for each participant. Word recognition thresholds corresponded with the VA level expected from normally sighted participants, and even a VA significantly higher than expected from normally sighted individuals for a few participants. Conclusions: Spatial resolution can be evaluated based on high-level stimuli encountered in day-to-day life, such as faces or written words with sweep visual evoked potentials.

A shared numerical magnitude representation evidenced by the distance effect in frequency-tagging EEG.

Humans can effortlessly abstract numerical information from various codes and contexts. However, whether the access to the underlying magnitude information relies on common or distinct brain representations remains highly debated. Here, we recorded electrophysiological responses to periodic variation of numerosity (every five items) occurring in rapid streams of numbers presented at 6 Hz in randomly varying codes-Arabic digits, number words, canonical dot patterns and finger configurations. Results demonstrated that numerical information was abstracted and generalized over the different representation codes by revealing clear discrimination responses (at 1.2 Hz) of the deviant numerosity from the base numerosity, recorded over parieto-occipital electrodes. Crucially, and supporting the claim that discrimination responses reflected magnitude processing, the presentation of a deviant numerosity distant from the base (e.g., base "2" and deviant "8") elicited larger right-hemispheric responses than the presentation of a close deviant numerosity (e.g., base "2" and deviant "3"). This finding nicely represents the neural signature of the distance effect, an interpretation further reinforced by the clear correlation with individuals' behavioral performance in an independent numerical comparison task. Our results therefore provide for the first time unambiguously a reliable and specific neural marker of a magnitude representation that is shared among several numerical codes.

Is human face recognition lateralized to the right hemisphere due to neural competition with left-lateralized visual word recognition? A critical review.

The right hemispheric lateralization of face recognition, which is well documented and appears to be specific to the human species, remains a scientific mystery. According to a long-standing view, the evolution of language, which is typically substantiated in the left hemisphere, competes with the cortical space in that hemisphere available for visuospatial processes, including face recognition. Over the last decade, a specific hypothesis derived from this view according to which neural competition in the left ventral occipito-temporal cortex with selective representations of letter strings causes right hemispheric lateralization of face recognition, has generated considerable interest and research in the scientific community. Here, a systematic review of studies performed in various populations (infants, children, literate and illiterate adults, left-handed adults) and methodologies (behavior, lesion studies, (intra)electroencephalography, neuroimaging) offers little if any support for this reading lateralized neural competition hypothesis. Specifically, right-lateralized face-selective neural activity already emerges at a few months of age, well before reading acquisition. Moreover, consistent evidence of face recognition performance and its right hemispheric lateralization being modulated by literacy level during development or at adulthood is lacking. Given the absence of solid alternative hypotheses and the key role of neural competition in the sensory-motor cortices for selectivity of representations, learning, and plasticity, a revised language-related neural competition hypothesis for the right hemispheric lateralization of face recognition should be further explored in future research, albeit with substantial conceptual clarification and advances in methodological rigor.

Dissociated face- and word-selective intracerebral responses in the human ventral occipito-temporal cortex.

The extent to which faces and written words share neural circuitry in the human brain is actively debated. Here, we compare face-selective and word-selective responses in a large group of patients (N = 37) implanted with intracerebral electrodes in the ventral occipito-temporal cortex (VOTC). Both face-selective (i.e., significantly different responses to faces vs. non-face visual objects) and word-selective (i.e., significantly different responses to words vs. pseudofonts) neural activity is isolated with frequency-tagging. Critically, this sensitive approach allows to objectively quantify category-selective neural responses and disentangle them from general visual responses. About 70% of significant electrode contacts show either face-selectivity or word-selectivity only, with the expected right and left hemispheric dominance, respectively. Spatial dissociations are also found within core regions of face and word processing, with a medio-lateral dissociation in the fusiform gyrus (FG) and surrounding sulci, respectively. In the 30% of overlapping face- and word-selective contacts across the VOTC or in the FG and surrounding sulci, between-category-selective amplitudes (faces vs. words) show no-to-weak correlations, despite strong correlations in both the within-category-selective amplitudes (face-face, word-word) and the general visual responses to words and faces. Overall, these observations support the view that category-selective circuitry for faces and written words is largely dissociated in the human adult VOTC.

Canonical representations of fingers and dots trigger an automatic activation of number semantics: an EEG study on 10-year-old children.

Over the course of development, children must learn to map a non-symbolic representation of magnitude to a more precise symbolic system. There is solid evidence that finger and dot representations can facilitate or even predict the acquisition of this mapping skill. While several behavioral studies demonstrated that canonical representations of fingers and dots automatically activate number semantics, no study so far has investigated their cerebral basis. To examine these questions, 10-year-old children were presented a behavioral naming task and a Fast Periodic Visual Stimulation EEG paradigm. In the behavioral task, children had to name as fast and as accurately as possible the numbers of dots and fingers presented in canonical and non-canonical configurations. In the EEG experiment, one category of stimuli (e.g., canonical representation of fingers or dots) was periodically inserted (1/5) in streams of another category (e.g., non-canonical representation of fingers or dots) presented at a fast rate (4 Hz). Results demonstrated an automatic access to number semantics and bilateral categorical responses at 4 Hz/5 for canonical representations of fingers and dots. Some differences between finger and dot configuration's processing were nevertheless observed and are discussed in light of an effortful-automatic continuum hypothesis.

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.