Christopher S.Y. Benwell

On the neural origin of pseudoneglect: EEG-correlates of shifts in line bisection performance with manipulation of line length.

Healthy participants tend to show systematic biases in spatial attention, usually to the left. However, these biases can shift rightward as a result of a number of experimental manipulations. Using electroencephalography (EEG) and a computerized line bisection task, here we investigated for the first time the neural correlates of changes in spatial attention bias induced by line-length (the so-called line-length effect). In accordance with previous studies, an overall systematic left bias (pseudoneglect) was present during long line but not during short line bisection performance. This effect of line-length on behavioral bias was associated with stronger right parieto-occipital responses to long as compared to short lines in an early time window (100–200 ms) post-stimulus onset. This early differential activation to long as compared to short lines was task-independent (present even in a non-spatial control task not requiring line bisection), suggesting that it reflects a reflexive attentional response to long lines. This was corroborated by further analyses source-localizing the line-length effect to the right temporo-parietal junction (TPJ) and revealing a positive correlation between the strength of this effect and the magnitude by which long lines (relative to short lines) drive a behavioral left bias across individuals. Therefore, stimulus-driven left bisection bias was associated with increased right hemispheric engagement of areas of the ventral attention network. This further substantiates that this network plays a key role in the genesis of spatial bias, and suggests that post-stimulus TPJ-activity at early information processing stages (around the latency of the N1 component) contributes to the left bias.

A rightward shift in the visuospatial attention vector with healthy aging

The study of lateralized visuospatial attention bias in non-clinical samples has revealed a systematic group-level leftward bias (pseudoneglect), possibly as a consequence of right hemisphere (RH) dominance for visuospatial attention. Pseudoneglect appears to be modulated by age, with a reduced or even reversed bias typically present in elderly participants. It has been suggested that this shift in bias may arise due to disproportionate aging of the RH and/or an increase in complementary functional recruitment of the left hemisphere (LH) for visuospatial processing. In this study, we report rightward shifts in subjective midpoint judgment relative to healthy young participants whilst elderly participants performed a computerized version of the landmark task (in which they had to judge whether a transection mark appeared closer to the right or left end of a line) on three different line lengths. This manipulation of stimulus properties led to a similar behavioral pattern in both the young and the elderly: a rightward shift in subjective midpoint with decreasing line length, which even resulted in a systematic rightward bias in elderly participants for the shortest line length (1.98° of visual angle, VA). Overall performance precision for the task was lower in the elderly participants regardless of line length, suggesting reduced landmark task discrimination sensitivity with healthy aging. This rightward shift in the attentional vector with healthy aging is likely to result from a reduction in RH resources/dominance for attentional processing in elderly participants. The significant rightward bias in the elderly for short lines may even suggest a reversal of hemisphere dominance in favor of the LH/right visual field under specific conditions.

Age-related reduction of hemispheric lateralisation for spatial attention: An EEG study

Abstract A group‐level visuospatial attention bias towards the left side of space (pseudoneglect) is consistently observed in young adults, which is likely to be a consequence of right parieto‐occipital dominance for spatial attention. Conversely, healthy older adults demonstrate a rightward shift of this behavioural bias, hinting that an age‐related reduction of lateralised neural activity may occur within visuospatial attention networks. We compared young (aged 18–25) and older (aged 60–80) adults on a computerised line bisection (landmark) task whilst recording event‐related potentials (ERPs). Full‐scalp cluster mass permutation tests identified a larger right parieto‐occipital response for long lines compared to short in young adults (confirming Benwell et al., 2014a) which was not present in the older group. To specifically investigate age‐related differences in hemispheric lateralisation, cluster mass permutation tests were then performed on a lateralised EEG dataset (RH‐LH electrodes). A period of right lateralisation was identified in response to long lines in young adults, which was not present for short lines. No lateralised clusters were present for either long or short lines in older adults. Additionally, a reduced P300 component amplitude was observed for older adults relative to young. We therefore report here, for the first time, an age‐related and stimulus‐driven reduction of right hemispheric control of spatial attention in older adults. Future studies will need to determine whether this is representative of the normal aging process or an early indicator of neurodegeneration. HighlightsNeural correlates of the landmark task compared in young vs older adults using EEG.Small leftward behavioral bias (pseudoneglect) at baseline for long lines in young.Right‐lateralised cluster of early EEG activity identified for long lines in young.No lateralised activity for either line length in older adults.Age‐related reduction of P300 component amplitude.

Inconsistent Effects of Parietal α-tACS on Pseudoneglect across Two Experiments: A Failed Internal Replication

Transcranial electrical stimulation (tES) is being investigated as an experimental and clinical interventional technique in human participants. While promising, important limitations have been identified, including weak effect sizes and high inter- and intra-individual variability of outcomes. Here, we compared two “inhibitory” tES-techniques with supposedly different mechanisms of action as to their effects on performance in a visuospatial attention task, and report on a direct replication attempt. In two experiments, 2 × 20 healthy participants underwent tES in three separate sessions testing different protocols (10 min stimulation each) with a montage targeting right parietal cortex (right parietal–left frontal, electrode-sizes: 3cm × 3cm–7 cm × 5 cm), while performing a perceptual line bisection (landmark) task. The tES-protocols were compared as to their ability to modulate pseudoneglect (thought to be under right hemispheric control). In experiment 1, sham-tES was compared to transcranial alternating current stimulation at alpha frequency (10 Hz; α-tACS) (expected to entrain “inhibitory” alpha oscillations) and to cathodal transcranial direct current stimulation (c-tDCS) (shown to suppress neuronal spiking activity). In experiment 2, we attempted to replicate the findings of experiment 1, and establish frequency-specificity by adding a 45 Hz-tACS condition to α-tACS and sham. In experiment 1, right parietal α-tACS led to the expected changes in spatial attention bias, namely a rightward shift in subjective midpoint estimation (relative to sham). However, this was not confirmed in experiment 2 and in the complete sample. Right parietal c-tDCS and 45 Hz-tACS had no effect. These results highlight the importance of replication studies, adequate statistical power and optimizing tES-interventions for establishing the robustness and reliability of electrical stimulation effects, and best practice.

Trial‐by‐trial co‐variation of pre‐stimulus EEG alpha power and visuospatial bias reflects a mixture of stochastic and deterministic effects

Human perception of perithreshold stimuli critically depends on oscillatory EEG activity prior to stimulus onset. However, it remains unclear exactly which aspects of perception are shaped by this pre‐stimulus activity and what role stochastic (trial‐by‐trial) variability plays in driving these relationships. We employed a novel jackknife approach to link single‐trial variability in oscillatory activity to psychometric measures from a task that requires judgement of the relative length of two line segments (the landmark task). The results provide evidence that pre‐stimulus alpha fluctuations influence perceptual bias. Importantly, a mediation analysis showed that this relationship is partially driven by long‐term (deterministic) alpha changes over time, highlighting the need to account for sources of trial‐by‐trial variability when interpreting EEG predictors of perception. These results provide fundamental insight into the nature of the effects of ongoing oscillatory activity on perception. The jackknife approach we implemented may serve to identify and investigate neural signatures of perceptual relevance in more detail.

EEG Microstate Correlates of Fluid Intelligence and Response to Cognitive Training

The neurobiological correlates of human fluid intelligence (Gf) remain elusive. Here, we demonstrate that spatiotemporal dynamics of EEG activity correlate with baseline measures of Gf and with its modulation by cognitive training. EEG dynamics were assessed in 74 healthy participants by examination of fast-changing, recurring, topographically-defined electric patterns termed “microstates”, which characterize the electrophysiological activity of distributed cortical networks. We find that the frequency of appearance of specific brain topographies, spatially associated with visual (microstate B) and executive control (microstate C) networks, respectively, is inversely related to Gf scores. Moreover, changes in Gf scores with cognitive training are inversely correlated with changes in microstate properties, indicating that the changes in brain network dynamics are behaviorally relevant. Finally, we find that cognitive training that increases Gf scores results in a posterior shift in the topography of microstate C. These results highlight the role of fast-changing brain electrical states in individual variability in Gf and in the response to cognitive training.

Efficacy of home-based visuomotor feedback training in stroke patients with chronic hemispatial neglect

ABSTRACT Hemispatial neglect is a severe cognitive condition frequently observed after a stroke, associated with unawareness of one side of space, disability and poor long-term outcome. Visuomotor feedback training (VFT) is a neglect rehabilitation technique that involves a simple, inexpensive and feasible training of grasping-to-lift rods at the centre. We compared the immediate and long-term effects of VFT vs. a control training when delivered in a home-based setting. Twenty participants were randomly allocated to an intervention (who received VFT) or a control group (n = 10 each). Training was delivered for two sessions by an experimenter and then patients self-administered it for 10 sessions over two weeks. Outcome measures included the Behavioural Inattention Test (BIT), line bisection, Balloons Test, Landmark task, room description task, subjective straight-ahead pointing task and the Stroke Impact Scale. The measures were obtained before, immediately after the training sessions and after four-months post-training. Significantly greater short and long-term improvements were obtained after VFT when compared to control training in line bisection, BIT and spatial bias in cancellation. VFT also produced improvements on activities of daily living. We conclude that VFT is a feasible, effective, home-based rehabilitation method for neglect patients that warrants further investigation with well-designed randomised controlled trials on a large sample of patients.

No changes in parieto-occipital alpha during neural phase locking to visual quasi-periodic theta-, alpha-, and beta-band stimulation

Recent studies have probed the role of the parieto‐occipital alpha rhythm (8–12 Hz) in human visual perception through attempts to drive its neural generators. To that end, paradigms have used high‐intensity strictly‐periodic visual stimulation that created strong predictions about future stimulus occurrences and repeatedly demonstrated perceptual consequences in line with an entrainment of parieto‐occipital alpha. Our study, in turn, examined the case of alpha entrainment by non‐predictive low‐intensity quasi‐periodic visual stimulation within theta‐ (4–7 Hz), alpha‐ (8–13 Hz), and beta (14–20 Hz) frequency bands, i.e., a class of stimuli that resemble the temporal characteristics of naturally occurring visual input more closely. We have previously reported substantial neural phase‐locking in EEG recording during all three stimulation conditions. Here, we studied to what extent this phase‐locking reflected an entrainment of intrinsic alpha rhythms in the same dataset. Specifically, we tested whether quasi‐periodic visual stimulation affected several properties of parieto‐occipital alpha generators. Speaking against an entrainment of intrinsic alpha rhythms by non‐predictive low‐intensity quasi‐periodic visual stimulation, we found none of these properties to show differences between stimulation frequency bands. In particular, alpha band generators did not show increased sensitivity to alpha band stimulation and Bayesian inference corroborated evidence against an influence of stimulation frequency. Our results set boundary conditions for when and how to expect effects of entrainment of alpha generators and suggest that the parieto‐occipital alpha rhythm may be more inert to external influences than previously thought.

Prestimulus EEG power predicts conscious awareness but not objective visual performance

Abstract Prestimulus oscillatory neural activity has been linked to perceptual outcomes during performance of psychophysical detection and discrimination tasks. Specifically, the power and phase of low frequency oscillations have been found to predict whether an upcoming weak visual target will be detected or not. However, the mechanisms by which baseline oscillatory activity influences perception remain unclear. Recent studies suggest that the frequently reported negative relationship between α power and stimulus detection may be explained by changes in detection criterion (i.e., increased target present responses regardless of whether the target was present/absent) driven by the state of neural excitability, rather than changes in visual sensitivity (i.e., more veridical percepts). Here, we recorded EEG while human participants performed a luminance discrimination task on perithreshold stimuli in combination with single-trial ratings of perceptual awareness. Our aim was to investigate whether the power and/or phase of prestimulus oscillatory activity predict discrimination accuracy and/or perceptual awareness on a trial-by-trial basis. Prestimulus power (3–28 Hz) was inversely related to perceptual awareness ratings (i.e., higher ratings in states of low prestimulus power/high excitability) but did not predict discrimination accuracy. In contrast, prestimulus oscillatory phase did not predict awareness ratings or accuracy in any frequency band. These results provide evidence that prestimulus α power influences the level of subjective awareness of threshold visual stimuli but does not influence visual sensitivity when a decision has to be made regarding stimulus features. Hence, we find a clear dissociation between the influence of ongoing neural activity on conscious awareness and objective performance.

Frequency and power of human alpha oscillations drift systematically and independently with time-on-task

Oscillatory neural activity is a fundamental characteristic of the mammalian brain spanning multiple levels of spatial and temporal scale. Current theories of neural oscillations and analysis techniques employed to investigate their functional significance are based on an often implicit assumption: In the absence of experimental manipulation, the spectral content of any given EEG- or MEG-recorded neural oscillator remains approximately stationary over the course of a typical experimental session (~1 hour), spontaneously fluctuating only around its dominant frequency. Here, we examined this assumption for ongoing neural oscillations in the alpha-band (8:13 Hz). We found that alpha peak frequency systematically decreased over time, while alpha-power increased. Intriguingly, these systematic changes showed partial independence of each other: Statistical source separation (independent component analysis) revealed that while some alpha components displayed concomitant power increases and peak frequency decreases, other components showed either unique power increases or frequency decreases. Interestingly, we also found these components to differ in frequency. Components that showed mixed frequency/power changes oscillated primarily in the lower alpha-band (~8-10Hz), while components with unique changes oscillated primarily in the higher alpha-band (~9-13Hz). Our findings provide novel clues on the time-varying intrinsic properties of large-scale neural networks as measured by M/EEG, with implications for the analysis and interpretation of studies that aim at identifying functionally relevant oscillatory networks or at driving them through external stimulation.