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.
We employed a jackknife method to link pre‐stimulus neural oscillations to psychometric measures during line bisection performance. The analysis identified both spontaneous trial‐by‐trial (stochastic) and systematic long‐term (deterministic) predictors of visuospatial bias, but no predictors of visual sensitivity. The approach we implemented helps to further characterize the influence of ongoing neural activity on perception and cognition.
Systematic biases in spatial attention are a common finding. In the general population, a systematic leftward bias is typically observed (pseudoneglect), possibly as a consequence of right hemisphere ...dominance for visuospatial attention. However, this leftward bias can cross-over to a systematic rightward bias with changes in stimulus and state factors (such as line length and arousal). The processes governing these changes are still unknown. Here we tested models of spatial attention as to their ability to account for these effects. To this end, we experimentally manipulated both stimulus and state factors, while healthy participants performed a computerized version of a landmark task. State was manipulated by time-on-task (>1 h) leading to increased fatigue and a reliable left- to rightward shift in spatial bias. Stimulus was manipulated by presenting either long or short lines which was associated with a shift of subjective midpoint from a reliable leftward bias for long to a more rightward bias for short lines. Importantly, we found time-on-task and line length effects to be additive suggesting a common denominator for line bisection across all conditions, which is in disagreement with models that assume that bisection decisions in long and short lines are governed by distinct processes (Magnitude estimation vs Global/local distinction). Our findings emphasize the dynamic rather than static nature of spatial biases in midline judgement. They are best captured by theories of spatial attention positing that spatial bias is flexibly modulated, and subject to inter-hemispheric balance which can change over time or conditions to accommodate task demands or reflect fatigue.
Pre‐stimulus electroencephalogram (EEG) oscillations, especially in the alpha range (8–13 Hz), can affect the sensitivity to temporal lags between modalities in multisensory perception. The effects ...of alpha power are often explained in terms of alpha's inhibitory functions, whereas effects of alpha frequency have bolstered theories of discrete perceptual cycles, where the length of a cycle, or window of integration, is determined by alpha frequency. Such studies typically employ visual detection paradigms with near‐threshold or even illusory stimuli. It is unclear whether such results generalize to above‐threshold stimuli. Here, we recorded EEG, while measuring temporal discrimination sensitivity in a temporal‐order judgement task using above‐threshold auditory and visual stimuli. We tested whether the power and instantaneous frequency of pre‐stimulus oscillations predict audiovisual temporal discrimination sensitivity on a trial‐by‐trial basis. By applying a jackknife procedure to link single‐trial pre‐stimulus oscillatory power and instantaneous frequency to psychometric measures, we identified a posterior cluster where lower alpha power was associated with higher temporal sensitivity of audiovisual discrimination. No statistically significant relationship between instantaneous alpha frequency and temporal sensitivity was found. These results suggest that temporal sensitivity for above‐threshold multisensory stimuli fluctuates from moment to moment and is indexed by modulations in alpha power.
In an audiovisual temporal‐order judgement task, we employed a novel jackknife procedure to quantify temporal sensitivity derived from a psychometric function at the single‐trial level. We found that higher pre‐stimulus alpha power predicted a lower temporal sensitivity, while instantaneous alpha frequency was not predictive of temporal sensitivity. Individual peak‐alpha frequency did not correlate significantly with temporal sensitivity across participants nor did individual peak‐alpha power.
Asymmetry in human spatial attention has long been documented. In the general population the majority of individuals tend to misbisect horizontal lines to the left of veridical centre. Nonetheless in ...virtually all previously reported studies on healthy participants, there have been subsets of people displaying rightward biases.
In this study, we report differential time-on task effects depending on participants' initial pseudoneglect bias: participants with an initial left bias in a landmark task (in which they had to judge whether a transection mark appeared closer to the right or left end of a line) showed a significant rightward shift over the course of the experimental session, whereas participants with an initial right bias shifted leftwards.
We argue that these differences in initial biases as well as the differential shifts with time-on task reflect genuine observer subtypes displaying diverging behavioural patterns. These observer subtypes could be driven by differences in brain organisation and/or lateralisation such as varying anatomical pathway asymmetries (Thiebaut de Schotten et al., 2011).
•Spatial attention biases have been found in both the diseased and healthy brain.•Healthy participants show left attentional biases with a minority displaying right biases.•We demonstrate that these differences in initial bias lead to differential time on-task effects.•We argue for genuine observer subtypes driven by differences in brain organisation.
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.
Using EEG we tested whether intrinsic alpha rhythms entrain to quasi‐periodic low‐intensity visual stimulation. Our results speak against a stimulus‐driven alpha entrainment when the quasiperiodicity of the stimulation is task‐irrelevant.
Rhythmic neural activity has been proposed to play a fundamental role in cognition. Both healthy and pathological aging are characterized by frequency-specific changes in oscillatory activity. ...However, the cognitive relevance of these changes across the spectrum from normal to pathological aging remains unknown. We examined electroencephalography (EEG) correlates of cognitive function in healthy aging and 2 of the most prominent and debilitating age-related disorders: type 2 diabetes mellitus (T2DM) and Alzheimer's disease (AD). Relative to healthy controls (HC), patients with AD were impaired on nearly every cognitive measure, whereas patients with T2DM performed worse mainly on learning and memory tests. A continuum of alterations in resting-state EEG was associated with pathological aging, generally characterized by reduced alpha (α) and beta (β) power (AD < T2DM < HC) and increased delta (δ) and theta (θ) power (AD > T2DM > HC), with some variations across different brain regions. There were also reductions in the frequency and power density of the posterior dominant rhythm in AD. The ratio of (α + β)/(δ + θ) was specifically associated with cognitive function in a domain- and diagnosis-specific manner. The results thus captured both similarities and differences in the pathophysiology of cerebral oscillations in T2DM and AD. Overall, pathological brain aging is marked by a shift in oscillatory power from higher to lower frequencies, which can be captured by a single cognitively relevant measure of the ratio of (α + β) over (δ + θ) power.
•Cognitive decline is widespread in Alzheimer's disease and selective in type 2 diabetes mellitus.•High delta (δ)/theta (θ) + low alpha (α)/beta (β) EEG power index pathological aging.•The α + β/δ + θ ratio predicts cognitive function, particularly symptom severity in Alzheimer's disease.
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.
Electroencephalography (EEG) has shown potential for identifying early-stage biomarkers of neurocognitive dysfunction associated with dementia due to Alzheimer's disease (AD). A large body of ...evidence shows that, compared to healthy controls (HC), AD is associated with power increases in lower EEG frequencies (delta and theta) and decreases in higher frequencies (alpha and beta), together with slowing of the peak alpha frequency. However, the pathophysiological processes underlying these changes remain unclear. For instance, recent studies have shown that apparent shifts in EEG power from high to low frequencies can be driven either by frequency specific periodic power changes or rather by non-oscillatory (aperiodic) changes in the underlying 1/f slope of the power spectrum. Hence, to clarify the mechanism(s) underlying the EEG alterations associated with AD, it is necessary to account for both periodic and aperiodic characteristics of the EEG signal. Across two independent datasets, we examined whether resting-state EEG changes linked to AD reflect true oscillatory (periodic) changes, changes in the aperiodic (non-oscillatory) signal, or a combination of both. We found strong evidence that the alterations are purely periodic in nature, with decreases in oscillatory power at alpha and beta frequencies (AD < HC) leading to lower (alpha + beta) / (delta + theta) power ratios in AD. Aperiodic EEG features did not differ between AD and HC. By replicating the findings in two cohorts, we provide robust evidence for purely oscillatory pathophysiology in AD and against aperiodic EEG changes. We therefore clarify the alterations underlying the neural dynamics in AD and emphasize the robustness of oscillatory AD signatures, which may further be used as potential prognostic or interventional targets in future clinical investigations.
Up to 80% of people who experience a right-hemisphere stroke suffer from hemispatial neglect. This syndrome is debilitating and impedes rehabilitation. We carried out a clinical feasibility trial of ...transcranial direct current stimulation (tDCS) and a behavioural rehabilitation programme, alone or in combination, in patients with neglect. Patients >4 weeks post right hemisphere stroke were randomized to 10 sessions of tDCS, 10 sessions of a behavioural intervention, combined intervention, or a control task. Primary outcomes were recruitment and retention rates, with secondary outcomes effect sizes on measures of neglect and quality of life, assessed directly after the interventions, and at 6 months follow up. Of 288 confirmed stroke cases referred (representing 7% of confirmed strokes), we randomized 8% (0.6% of stroke cases overall). The largest number of exclusions (91/288 (34%)) were due to medical comorbidities that prevented patients from undergoing 10 intervention sessions. We recruited 24 patients over 29 months, with 87% completing immediate post-intervention and 67% 6 month evaluations. We established poor feasibility of a clinical trial requiring repeated hospital-based tDCS within a UK hospital healthcare setting, either with or without behavioural training, over a sustained time period. Future trials should consider intensity, duration and location of tDCS neglect interventions.
Trial registration:
ClinicalTrials.gov identifier: NCT02401724
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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.
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