Sustained visuospatial attention enhances lateralized anticipatory ERP activity in sensory areas.

The existence of neural correlates of spatial attention is not limited to the reactive stage of stimulus processing: neural activities subtending spatial attention are deployed well ahead of stimulus onset. ERP evidence supporting this proactive (top-down) attentional control is based on trial-by-trial S1-S2 paradigms, where the onset of a directional cue (S1) indicates on which side attention must be directed to respond to an upcoming target stimulus (S2). Crucially, S1 onset trigger both attention and motor preparation, therefore, these paradigms are not ideal to demonstrate the effect of attention at preparatory stage of processing. To isolate top-down anticipatory attention, the present study used a sustained attention paradigm based on a steady cue that indicates the attended side constantly throughout an entire block of trials, without any onset of an attentional cue. The main result consists in the description of the attention effect on the visual negativity (vN) component, a growing neural activity starting before stimulus presentation in extrastriate visual areas. The vN was consistently lateralized in the hemisphere contralateral to the attended side, regardless of the hand to be used. At the opposite, the lateralized motor activity emerged long after, confirming that the hand-selection process followed the spatial attention orientation process. The present study confirms the anticipatory nature of the vN component and corroborate its role in terms of preparatory visuospatial attention.

Testing the Specificity of Predictors of Reading, Spelling and Maths: A New Model of the Association Among Learning Skills Based on Competence, Performance and Acquisition.

In a previous study (Zoccolotti et al., 2020) we examined reading, spelling, and maths skills in an unselected group of 129 Italian children attending fifth grade by testing various cognitive predictors; results showed a high degree of predictors' selectivity for each of these three behaviors. In the present study, we focused on the specificity of the predictors by performing cross-analyses on the same dataset; i.e., we predicted spelling and maths skills based on reading predictors, reading based on maths predictors and so on. Results indicated that some predictors, such as the Orthographic Decision and the Arithmetic Facts tests, predicted reading, spelling and maths skills in similar ways, while others predicted different behaviors but only for a specific parameter, such as fluency but not accuracy (as in the case of RAN), and still others were specific for a single behavior (e.g., Visual-auditory Pseudo-word Matching test predicted only spelling skills). To interpret these results, we propose a novel model of learning skills separately considering factors in terms of competence, performance and acquisition (automatization). Reading, spelling and calculation skills would depend on the development of discrete and different abstract competences (accounting for the partial dissociations among learning disorders reported in the literature). By contrast, overlap among behaviors would be accounted for by defective acquisition in automatized responses to individual "instances"; this latter skill is item specific but domain independent. Finally, performance factors implied in task's characteristics (such as time pressure) may contribute to the partial association among learning skills. It is proposed that this new model may provide a useful base for interpreting the diffuse presence of comorbidities among learning disorders.

Preparatory ERPs in visual, auditory, and somatosensory discriminative motor tasks.

Previous event-related potential (ERP) studies mainly from the present research group showed a novel component, that is, the prefrontal negativity (pN), recorded in visual-motor discriminative tasks during the pre-stimulus phase. This component is concomitant to activity related to motor preparation, that is, the Bereitschaftspotential (BP). The pN component has been reported in experiments based on the visual modality only; for other modalities (acoustic and/or somatosensory) the presence of the pN warrants further investigation. This study represents a first step toward this direction; indeed, we aimed at describing the pN and the BP components in discriminative response tasks (DRTs) for three sensory modalities. In experiment 1 ERPs were recorded in 29 adults in visual and auditory DRT; an additional group of 15 adults participated to a somatosensory DRT (experiment 2). In line with previous results both the pN and the BP were clearly detectable in the visual modality. In the auditory modality the prefrontal pN was not detectable directly; however, the pN could be derived by subtraction of separate EEG traces recorded in a "passive" version of the same auditory task, in which motor responses were not required. In the somatosensory modality both the pN and the BP were detectable, although with lower amplitudes with respect to other two sensory modalities. Overall, regardless of the sensory modality, anticipatory task-related pN and BP components could be detected (or derived by subtraction) over both the prefrontal and motor cortices. These results support the view that anticipatory processes share common components among sensory modalities.

Predicting individual differences in reading, spelling and maths in a sample of typically developing children: A study in the perspective of comorbidity.

We examined reading, spelling, and mathematical skills in an unselected group of 129 Italian fifth graders by testing various cognitive predictors for each behaviour. As dependent variables, we measured performance in behaviours with a clear functional value in everyday life, such as reading a text, spelling under dictation and doing mental and written computations. As predictors, we selected cognitive dimensions having an explicit relation with the target behaviour (called proximal predictors), and prepared various tests in order to select which task had the best predictive power on each behaviour. The aim was to develop a model of proximal predictors of reading (speed and accuracy), spelling (accuracy) and maths (speed and accuracy) characterized by efficacy also in comparison to the prediction based on general cognitive factors (i.e., short-term memory, phonemic verbal fluency, visual perceptual speed, and non-verbal intelligence) and parsimony, pinpointing the role of both common and unique predictors as envisaged in the general perspective of co-morbidity. With one exception (reading accuracy), the proximal predictors models (based on communality analyses) explained a sizeable amount of variance, ranging from 27.5% in the case of calculation (accuracy) to 48.7% of reading (fluency). Models based on general cognitive factors also accounted for some variance (ranging from 6.5% in the case of spelling to 19.5% in the case of reading fluency) but this was appreciably less than that explained by models based on the hypothesized proximal predictors. In general, results confirmed the efficacy of proximal models in predicting reading, spelling and maths although they offered only limited support for common predictors across different learning skills; namely, performance in the Orthographic Decision test entered as a predictor of both reading and spelling indicating that a single orthographic lexicon may account for performance in reading and spelling. Possible lines of research to expand on this approach are illustrated.

Prompting future events: Effects of temporal cueing and time on task on brain preparation to action.

Prediction about event timing plays a leading role in organizing and optimizing behavior. We recorded anticipatory brain activities and evaluated whether temporal orienting processes are reflected by the novel prefrontal negative (pN) component, as already shown for the contingent negative variation (CNV). Fourteen young healthy participants underwent EEG and fMRI recordings in separate sessions; they were asked to perform a Go/No-Go task in which temporal orienting was manipulated: the external condition (a visual display indicating the time of stimulus onset) and the internal condition (time information not provided). In both conditions, the source of the pN was localized in the pars opercularis of the iFg; the source of the CNV was localized in the supplementary motor area and cingulate motor area, as expected. Anticipatory activity was also found in the occipital-parietal cortex. Time on task EEG analysis showed a marked learning effect in the internal condition, while the effect was minor in the external condition. In fMRI, the two conditions had a similar pattern; similarities and differences of results obtained with the two techniques are discussed. Overall, data are consistent with the view that the pN reflects a proactive cognitive control, including temporal orienting.

Proactive Cortical Control in Spinal Cord Injury Subjects with Paraplegia.

Preparatory cortical activities were investigated in subjects with paraplegia attributed to spinal cord injury (SCI). Electroencephalogram (EEG) and behavioral data were recorded simultaneously in a visual-motor discrimination go/no-go task performed with the right upper limb. Eighteen SCI subjects participated to one, two, or three experimental sessions (Go/No-Go task), according to their availability and willingness to participate. To evaluate the effects of SCI on cortical activities as a function of time, we considered three SCI groups (9 individuals each), based on different time from the injury onset (acute, 1-2 months; subacute, 3-5 months; chronic, 6-9 months), and a control group of 9 healthy participants matched for age and sex. Results indicate that response time (RT) was slower and percentage of omissions was higher in SCI subjects than healthy, independently from time from lesion (TFL). Also, the proactive motor preparation, indexed by the Bereitschafts potential (BP), and the pre-frontal cognitive control, indexed by the pre-frontal negativity component, showed reduced amplitude in SCI subjects, independently from TFL. Conversely, TFL effect was observed in the BP topography, which showed a more posterior focus in subacute and chronic groups than healthies. Interestingly, despite this posteriorization, BP amplitudes maintained the well-known correlation with RTs. Overall, SCI affects cortical reorganization independently from TFL, regarding proactive activities for action inhibition and reaction time; conversely, a TFL effect was observed in the topography changes related to the cortical areas involved in proactive motor activity. Present data are in line with growing evidence of brain changes after SCI, in particular focusing on cognitive effects and evidencing possible functional mechanisms related to motor and cognitive readiness processing, relevant for SCI rehabilitation programs.

Perceptual load in decision making: The role of anterior insula and visual areas. An ERP study.

The present study aimed at describing the effects of perceptual load on neurocognitive processes of decision-making. To this aim, we used a visual-motor discriminative task in which pairs of stimuli were assigned to either target or non-target categories. For each category, stimulus configuration was defined as simple or complex according to orientation and arrangement of the constituent segments. Analyses of prefrontal ERPs revealed that the pP1 component (at 180 ms) was larger for complex stimuli than simple for both categories, and the same result was found for the pP2 component (at 320 ms). Occipital ERPs revealed effects of perceptual load on the N1 component, but not on the mainly exogenous P1 component, indicating that amplitude modulations of prefrontal ERPs were not due to physical difference between simple and complex stimuli. Based on the recent literature, we discussed the pP1 activity as reflecting a process of sensory-motor integration, and the pP2 as a stimulus-response mapping process resulting from two gradients of activity: one category-based (larger for target than non-target stimuli), the other decision effort-based (enhanced when categorization implied a greater attentional load). Previous ERP-fMRI studies and present source analysis support the view that prefrontal ERPs were generated mainly by activity in the anterior Insula.

Electrophysiological evidence of sustained spatial attention effects over anterior cortex: Possible contribution of the anterior insula.

Spatial attention can improve performance in terms of speed and accuracy; this advantage may be mediated by brain processes at both poststimulus (reactive) and prestimulus (proactive) stages. Here, we studied how visuospatial attention affects both proactive and reactive brain functions using event-related potentials (ERPs). At reactive stage, effects of attention on parietal-occipital components are well documented; little data are available on anterior components. Seventeen participants performed simple and discriminative response tasks, while voluntarily and steadily attending either the left or right visual hemifield throughout one block. Response speed was faster for the attended side. At ERP level, attending to one hemifield did not produce lateralization of proactive components-that is, the BP and the pN. As for poststimulus components, we confirmed the well-known amplitude effects on the P1, N1, and P3. More interesting are results for the prefrontal components previously neglected in tasks modulating spatial attention. Previous studies suggest that these components reflect perceptual and sensory-motor awareness (pN1 and pP1 components), and stimulus-response mapping (pP2 component) associated to anterior insular activity. Spatial attention enhanced the pN1 and the pP1 amplitude but had no effect on the pP2. Overall, results extend knowledge on spatial attention, showing that sustained spatial attention affects the activity of anterior areas, such as the anterior insula, in addition to the known influence on occipital-parietal areas. Top-down spatial attention is likely mediated by increased sensory and sensory-motor awareness for attended events; this effect is evident in reactive, not proactive, brain activity.

Awareness of perception and sensory-motor integration: ERPs from the anterior insula.

The present work follows recent evidences of studies showing that visual stimuli evoke two early prefrontal event-related potentials (ERP) concomitant to the canonical occipital activities, but originating within the anterior insula (the pN1 and the pP1 components). To clarify the exogenous/endogenous nature of these components, we performed two experiments in which stimulus physical features (Experiment 1) and motor demands of the task (Experiment 2) were considered. In a simple response task (SRT), low-visibility stimuli evoked larger pN1 over the prefrontal areas (Experiment 1) with respect to high-visibility stimuli; in contrast, the occipital P1 component (concomitant to the pN1) had reduced amplitude in the low-visibility condition as expected. Furthermore, the latency of the P1, pN1 and pP1 was slower in the low-visibility condition (from 8 to 18 ms), and the motor response was slowed down as well (on average 14 ms). Pre-stimulus analysis showed that low-visibility stimuli were preceded by greater motor readiness. On the other hand, Experiment 2 showed that, compared with the SRT, the request to passively view the same stimuli was associated with smaller pP1. ERP source analysis confirmed the anterior insula source of the prefrontal ERPs; we interpreted these activities as the correlate of two top-down perceptual processing: the sensory awareness (the pN1) and the awareness of the sensory-motor integration (the pP1), associated with the subjective experience of the visual perception and the conscious experience of the sensory-motor coupling, respectively.

Brain waves from an "isolated" cortex: contribution of the anterior insula to cognitive functions.

Using two independent electrical neuroimaging techniques (BESA and sLORETA), we tested a fMRI-seeded source modeling indicating that in visual discriminative tasks the anterior insula (aIns) participates in the generation of three prefrontal ERP components: the pN1 (at 115 ms), the pP1 (at 170 ms), and the pP2 (at 300 ms). This latter component represented the focus of the present study. Results showed that the pP2 had different activation profiles across hemispheres. The left aIns activity peaked at 420 ms (30 ms before the response) for both Go and No-go trials, that is independently from the ultimate choice (response or inhibition). The right aIns activity started at about 250 ms and progressively increased for a time interval extending after the motor response; its amplitude was larger in case of Go than No-go stimuli. We suggest that the activation of the left aIns reflected the timing of the decision, and the right aIns the categorization and the performance monitoring processes. A control experiment requiring simple (not discriminative) motor response revealed that the pP2 and the aIns activity were nearly absent after the 250 ms; this result confirmed that the aIns activity at this stage is associated with the decisional processes, and not with the motor response per se. The present investigation shed new lights on the insular contribution to perceptual decision-making, and opens to the possibility of assessing the aIns activity via ERP analysis.

Reading and lexical-decision tasks generate different patterns of individual variability as a function of condition difficulty.

We reanalyzed previous experiments based on lexical-decision and reading-aloud tasks in children with dyslexia and control children and tested the prediction of the difference engine model (DEM) that mean condition reaction times (RTs) and standard deviations (SDs) would be linearly related (Myerson et al., 2003). Then we evaluated the slope and the intercept with the x-axis of these linear functions in comparison with previously reported values (i.e., slope of about 0.30 and intercept of about 300 ms). In the case of lexical decision, the parameters were close to these values; by contrast, in the case of reading aloud, a much steeper slope (0.66) and a greater intercept (482.6 ms) were found. Therefore, interindividual variability grows at a much faster rate as a function of condition difficulty for reading than for lexical-decision tasks (or for other tasks reported in the literature). According to the DEM, the slope of the regression that relates means and SDs indicates the degree of correlation among the durations of the stages of processing. We propose that the need for a close coupling between orthographic and phonological processing in reading is what drives the particularly strong relationship between performance and interindividual variability that we observed in reading tasks.

Slowing in reading and picture naming: the effects of aging and developmental dyslexia.

We examined the slowing in vocal reaction times shown by dyslexic (compared to control) children with that of older (compared to younger) adults using an approach focusing on the detection of global, non-task-specific components. To address this aim, data were analyzed with reference to the difference engine (DEM) and rate and amount (RAM) models. In Experiment 1, typically developing children, children with dyslexia (both attending sixth grade), younger adults and older adults read words and non-words and named pictures. In Experiment 2, word and picture conditions were presented to dyslexic and control children attending eighth grade. In both experiments, dyslexic children were delayed in reading conditions, while they were unimpaired in naming pictures (a finding which indicates spared access to the phonological lexicon). The reading difficulty was well accounted for by a single multiplicative factor while only the residual effect of length (but not frequency and lexicality) was present after controlling for over-additivity using a linear mixed effects model with random slopes on critical variables. Older adults were slower than younger adults across reading and naming conditions. This deficit was well described by a single multiplicative factor. Thus, while slowing of information processing is limited to orthographic stimuli in dyslexic children, it cuts across verbal tasks in older adults. Overall, speed differences in groups such as dyslexic children and older adults can be effectively described with reference to deficits in domains encompassing a variety of experimental conditions rather than deficits in single specific task/conditions. The DEM and RAM prove effective in teasing out global vs. specific components of performance.

Missing the Target: the Neural Processing Underlying the Omission Error.

The omissions are infrequent errors consisting in missing responses to the target stimuli. This is the first study aimed at investigating the brain activities associated with omissions in a decision-making task. We recorded event-related potentials (ERPs) in 12 subjects which reported a suitable number of omissions in a visual go/no-go task. We investigated both the pre- and post-stimulus brain activities associated with correct and omitted trials. The electrical neuroimaging technique (BESA) was adopted to extract the anterior insula (aIns) activity associated with the prefrontal P2 component (pP2) peaking about 300 ms after the stimulus and reflecting the stimulus-response mapping process. We found that omissions were predicted by a delayed onset (about half a second) of two pre-stimulus components, i.e. the prefrontal negativity (pN) and the Bereitschaftspotential (BP) associated with the top-down control and the motor preparation, respectively. Further, at the post-stimulus stage the omission trials were characterized by the suppression of the pP2 (and the aIns activity as measured by BESA). No differences between omission and correct trials were detected at the level of the P1 and N1 visual components, as well as the P3. These findings would suggest that omissions are attentional lapsebased errors, as indicated by the delayed brain preparation before the stimulus onset. The reduced cortical activity during the preparation phase did not affect the visual processing; in contrast the stimulus categorization process at the level of the anterior insula did not start at all, resulting in the inability to reach a decision.

How the brain prevents a second error in a perceptual decision-making task.

In cognitive tasks, error commission is usually followed by a performance characterized by post-error slowing (PES) and post-error improvement of accuracy (PIA). Three theoretical accounts were hypothesized to support these post-error adjustments: the cognitive, the inhibitory, and the orienting account. The aim of the present ERP study was to investigate the neural processes associated with the second error prevention. To this aim, we focused on the preparatory brain activities in a large sample of subjects performing a Go/No-go task. The main results were the enhancement of the prefrontal negativity (pN) component -especially on the right hemisphere- and the reduction of the Bereitschaftspotential (BP) -especially on the left hemisphere- in the post-error trials. The ERP data suggested an increased top-down and inhibitory control, such as the reduced excitability of the premotor areas in the preparation of the trials following error commission. The results were discussed in light of the three theoretical accounts of the post-error adjustments. Additional control analyses supported the view that the adjustments-oriented components (the post-error pN and BP) are separated by the error-related potentials (Ne and Pe), even if all these activities represent a cascade of processes triggered by error-commission.

Rhythmic modulation of visual contrast discrimination triggered by action.

Recent evidence suggests that ongoing brain oscillations may be instrumental in binding and integrating multisensory signals. In this experiment, we investigated the temporal dynamics of visual-motor integration processes. We show that action modulates sensitivity to visual contrast discrimination in a rhythmic fashion at frequencies of about 5 Hz (in the theta range), for up to 1 s after execution of action. To understand the origin of the oscillations, we measured oscillations in contrast sensitivity at different levels of luminance, which is known to affect the endogenous brain rhythms, boosting the power of alpha-frequencies. We found that the frequency of oscillation in sensitivity increased at low luminance, probably reflecting the shift in mean endogenous brain rhythm towards higher frequencies. Importantly, both at high and at low luminance, contrast discrimination showed a rhythmic motor-induced suppression effect, with the suppression occurring earlier at low luminance. We suggest that oscillations play a key role in sensory-motor integration, and that the motor-induced suppression may reflect the first manifestation of a rhythmic oscillation.

Perceptual and Cognitive Factors Imposing "Speed Limits" on Reading Rate: A Study with the Rapid Serial Visual Presentation.

Adults read at high speed, but estimates of their reading rate vary greatly, i.e., from 100 to 1500 words per minute (wpm). This discrepancy is likely due to different recording methods and to the different perceptual and cognitive processes involved in specific test conditions. The present study investigated the origins of these notable differences in RSVP reading rate (RR). In six experiments we investigated the role of many different perceptual and cognitive variables. The presence of a mask caused a steep decline in reading rate, with an estimated masking cost of about 200 wpm. When the decoding process was isolated, RR approached values of 1200 wpm. When the number of stimuli exceeded the short-term memory span, RR decreased to 800 wpm. The semantic context contributed to reading speed only by a factor of 1.4. Finally, eye movements imposed an upper limit on RR (around 300 wpm). Overall, data indicate a speed limit of 300 wpm, which corresponds to the time needed for eye movement execution, i.e., the most time consuming mechanism. Results reconcile differences in reading rates reported by different laboratories and thus provide suggestions for targeting different components of reading rate.

Spatiotemporal brain mapping during preparation, perception, and action.

Deciding whether to act or not to act is a fundamental cognitive function. To avoid incorrect responses, both reactive and proactive modes of control have been postulated. Little is known, however, regarding the brain implementation of proactive mechanisms, which are deployed prior to an actual need to inhibit a response. Via a combination of electrophysiological and neuroimaging measures (recorded in 21 and 16 participants, respectively), we describe the brain localization and timing of neural activity that underlies the anticipatory proactive mechanism. From these results, we conclude that proactive control originates in the inferior Frontal gyrus, is established well before stimulus perception, and is released concomitantly with stimulus appearance. Stimulus perception triggers early activity in the anterior insula and intraparietal cortex contralateral to the responding hand; these areas likely mediate the transition from perception to action. The neural activities leading to the decision to act or not to act are described in the framework of a three-stage model that includes perception, action, and anticipatory functions taking place well before stimulus onset.

Discrete versus multiple word displays: a re-analysis of studies comparing dyslexic and typically developing children.

The study examines whether impairments in reading a text can be explained by a deficit in word decoding or an additional deficit in the processes governing the integration of reading subcomponents (including eye movement programming and pronunciation) should also be postulated. We report a re-analysis of data from eleven previous experiments conducted in our lab where the reading performance on single, discrete word displays as well multiple displays (texts, and in few cases also word lists) was investigated in groups of dyslexic children and typically developing readers. The analysis focuses on measures of time and not accuracy. Across experiments, dyslexic children are slower and more variable than typically developing readers in reading texts as well as vocal reaction time (RTs) to singly presented words; the dis-homogeneity in variability between groups points to the inappropriateness of standard measures of size effect (such as Cohen's d), and suggests the use of the ratio between groups' performance. The mean ratio for text reading is 1.95 across experiments. Mean ratio for vocal RTs for singly presented words is considerably smaller (1.52). Furthermore, this latter value is probably an overestimation as considering total reading times (i.e., a measure including also the pronunciation component) considerably reduces the group difference in vocal RTs (1.19 according to Martelli et al., 2014). The ratio difference between single and multiple displays does not depend upon the presence of a semantic context in the case of texts as large ratios are also observed with lists of unrelated words (though studies testing this aspect were few). We conclude that, if care is taken in using appropriate comparisons, the deficit in reading texts or lists of words is appreciably greater than that revealed with discrete word presentations. Thus, reading multiple stimuli present a specific, additional challenge to dyslexic children indicating that models of reading should incorporate this aspect.

Rhythmic oscillations of visual contrast sensitivity synchronized with action.

It is well known that the motor and the sensory systems structure sensory data collection and cooperate to achieve an efficient integration and exchange of information. Increasing evidence suggests that both motor and sensory functions are regulated by rhythmic processes reflecting alternating states of neuronal excitability, and these may be involved in mediating sensory-motor interactions. Here we show an oscillatory fluctuation in early visual processing time locked with the execution of voluntary action, and, crucially, even for visual stimuli irrelevant to the motor task. Human participants were asked to perform a reaching movement toward a display and judge the orientation of a Gabor patch, near contrast threshold, briefly presented at random times before and during the reaching movement. When the data are temporally aligned to the onset of movement, visual contrast sensitivity oscillates with periodicity within the theta band. Importantly, the oscillations emerge during the motor planning stage, ∼500 ms before movement onset. We suggest that brain oscillatory dynamics may mediate an automatic coupling between early motor planning and early visual processing, possibly instrumental in linking and closing up the visual-motor control loop.