Investigating predictive coding in younger and older children using MEG and a multi-feature auditory oddball paradigm.

There is mounting evidence for predictive coding theory from computational, neuroimaging, and psychological research. However, there remains a lack of research exploring how predictive brain function develops across childhood. To address this gap, we used pediatric magnetoencephalography to record the evoked magnetic fields of 18 younger children (M = 4.1 years) and 19 older children (M = 6.2 years) as they listened to a 12-min auditory oddball paradigm. For each child, we computed a mismatch field "MMF": an electrophysiological component that is widely interpreted as a neural signature of predictive coding. At the sensor level, the older children showed significantly larger MMF amplitudes relative to the younger children. At the source level, the older children showed a significantly larger MMF amplitude in the right inferior frontal gyrus relative to the younger children, P < 0.05. No differences were found in 2 other key regions (right primary auditory cortex and right superior temporal gyrus) thought to be involved in mismatch generation. These findings support the idea that predictive brain function develops during childhood, with increasing involvement of the frontal cortex in response to prediction errors. These findings contribute to a deeper understanding of the brain function underpinning child cognitive development.

Atypical neural responses to vocal anger in attention-deficit/hyperactivity disorder.

BACKGROUND: Deficits in facial emotion processing, reported in attention-deficit/hyperactivity disorder (ADHD), have been linked to both early perceptual and later attentional components of event-related potentials (ERPs). However, the neural underpinnings of vocal emotion processing deficits in ADHD have yet to be characterised. Here, we report the first ERP study of vocal affective prosody processing in ADHD. METHODS: Event-related potentials of 6-11-year-old children with ADHD (n = 25) and typically developing controls (n = 25) were recorded as they completed a task measuring recognition of vocal prosodic stimuli (angry, happy and neutral). Audiometric assessments were conducted to screen for hearing impairments. RESULTS: Children with ADHD were less accurate than controls at recognising vocal anger. Relative to controls, they displayed enhanced N100 and attenuated P300 components to vocal anger. The P300 effect was reduced, but remained significant, after controlling for N100 effects by rebaselining. Only the N100 effect was significant when children with ADHD and comorbid conduct disorder (n = 10) were excluded. CONCLUSION: This study provides the first evidence linking ADHD to atypical neural activity during the early perceptual stages of vocal anger processing. These effects may reflect preattentive hyper-vigilance to vocal anger in ADHD.

The reinforcing value of delay escape in attention deficit/hyperactivity disorder: An electrophysiological study.

The delay aversion hypothesis argues that the tendency for impulsive choice (preference for smaller sooner over larger later rewards) is motivated by the escape of negative affective states associated with delay. This model predicts that individuals with ADHD find the imposition of delay before an outcome or event especially aversive and its escape reinforcing. Consistent with this, fMRI studies show that ADHD is associated with amygdala hyper-sensitivity to cues of delay. However, evidence that delay escape is reinforcing is lacking. Here we extend fMRI research by using electrophysiological methods to study the reinforcing properties of delay-escape in ADHD. Thirty controls and 25 adolescents with ADHD aged 10-15 years performed the Escape Delay Incentive (EDI) task- in which pre-target cues indicated three conditions: i) CERTAIN DELAY: delay would follow a response irrespective of response speed ii) CONDITIONAL DELAY: delay would only follow if the response was too slow and iii) NO DELAY: delay would follow the response whatever the speed. We focused on the Contingent Negative Variation (CNV), a cue-evoked marker of motivated response preparation, across two time windows (CNV1 and CNV2). We took measures of parent, teacher and self-rated ADHD symptoms, task performance (RT) and self-rated delay aversion. We isolated CNV components and compared these between ADHD and controls. Adolescents with ADHD displayed a larger CNV2 to the CONDITIONAL DELAY than the CERTAIN DELAY cues compared to controls. However, this effect was not mirrored at the performance level and was unrelated to self-reported delay aversion. Our study provides the first ERP evidence that delay escape differentially reinforcers neural activation of attention preparation in ADHD cases. Future studies should examine the impact of varying cognitive load on task EDI performance.

The Moderating Effect of Self-Reported State and Trait Anxiety on the Late Positive Potential to Emotional Faces in 6-11-Year-Old Children.

Introduction: The emergence of anxiety during childhood is accompanied by the development of attentional biases to threat. However, the neural mechanisms underlying these biases are poorly understood. In addition, previous research has not examined whether state and trait anxiety are independently associated with threat-related biases. Methods: We compared ERP waveforms during the processing of emotional faces in a population sample of 58 6-11-year-olds who completed self-reported measures of trait and state anxiety and depression. Results: The results showed that the P1 was larger to angry than neutral faces in the left hemisphere, though early components (P1, N170) were not strongly associated with child anxiety or depression. In contrast, Late Positive Potential (LPP) amplitudes to angry (vs. neutral) faces were significantly and positively associated with symptoms of anxiety/depression. In addition, the difference between LPPs for angry (vs. neutral) faces was independently associated with state and trait anxiety symptoms. Discussion: The results showed that neural responses to facial emotion in children with elevated symptoms of anxiety and depression were most evident at later processing stages characterized as evaluative and effortful. The findings support cognitive models of threat perception in anxiety and indicate that trait elements of anxiety and more transitory fluctuations in anxious affect are important in understanding individual variation in the neural response to threat in late childhood.

An electrophysiological investigation of reinforcement effects in attention deficit/hyperactivity disorder: Dissociating cue sensitivity from down-stream effects on target engagement and performance.

OBJECTIVE: Neural hypo-sensitivity to cues predicting positive reinforcement has been observed in ADHD using the Monetary Incentive Delay (MID) task. Here we report the first study using an electrophysiological analogue of this task to distinguish between (i) cue related anticipation of reinforcement and downstream effects on (ii) target engagement and (iii) performance in a clinical sample of adolescents with ADHD and controls. METHODS: Thirty-one controls and 32 adolescents with ADHD aged 10-16 years performed the electrophysiological (e)-MID task - in which preparatory cues signal whether a response to an upcoming target will be reinforced or not - under three conditions; positive reinforcement, negative reinforcement (response cost) and no consequence (neutral). We extracted values for both cue-related potentials known to be, both, associated with response preparation and modulated by reinforcement (Cue P3 and Cue CNV) and target-related potentials (target P3) and compared these between ADHD and controls. RESULTS: ADHD and controls did not differ on cue-related components on neutral trials. Against expectation, adolescents with ADHD displayed Cue P3 and Cue CNV reinforcement-related enhancement (versus neutral trials) compared to controls. ADHD individuals displayed smaller target P3 amplitudes and slower and more variable performance - but effects were not modulated by reinforcement contingencies. When age, IQ and conduct problems were controlled effects were marginally significant but the pattern of results did not change. DISCUSSION: ADHD was associated with hypersensitivity to positive (and marginally negative) reinforcement reflected on components often thought to be associated with response preparation - however these did not translate into improved attention to targets. In the case of ADHD, upregulated CNV may be a specific marker of hyper-arousal rather than an enhancement of anticipatory attention to upcoming targets. Future studies should examine the effects of age, IQ and conduct problems on reinforcement sensitivity in ADHD.

Spontaneous activity in the waiting brain: a marker of impulsive choice in attention-deficit/hyperactivity disorder?

BACKGROUND: Spontaneous very low frequency oscillations (VLFO), seen in the resting brain, are attenuated when individuals are working on attention demanding tasks or waiting for rewards (Hsu et al., 2013). Individuals with attention-deficit/hyperactivity disorder (ADHD) display excess VLFO when working on attention tasks. They also have difficulty waiting for rewards. Here we examined the waiting brain signature in ADHD and its association with impulsive choice. METHODS: DC-EEG from 21 children with ADHD and 21 controls (9-15 years) were collected under four conditions: (i) resting; (ii) choosing to wait; (iii) being "forced" to wait; and (iv) working on a reaction time task. A questionnaire measured two components of impulsive choice. RESULTS: Significant VLFO reductions were observed in controls within anterior brain regions in both working and waiting conditions. Individuals with ADHD showed VLFO attenuation while working but to a reduced level and none at all when waiting. A closer inspection revealed an increase of VLFO activity in temporal regions during waiting. Excess VLFO activity during waiting was associated with parents' ratings of temporal discounting and delay aversion. CONCLUSIONS: The results highlight the potential role for waiting-related spontaneous neural activity in the pathophysiology of impulsive decision-making of ADHD.

The role of configurality in the Thatcher illusion: an ERP study.

The Thatcher illusion (Thompson in Perception, 9, 483-484, 1980) is often explained as resulting from recognising a distortion of configural information when 'Thatcherised' faces are upright but not when inverted. However, recent behavioural studies suggest that there is an absence of perceptual configurality in upright Thatcherised faces (Donnelly et al. in Attention, Perception & Psychophysics, 74, 1475-1487, 2012) and both perceptual and decisional sources of configurality in behavioural tasks with Thatcherised stimuli (Mestry, Menneer et al. in Frontiers in Psychology, 3, 456, 2012). To examine sources linked to the behavioural experience of the illusion, we studied inversion and Thatcherisation of faces (comparing across conditions in which no features, the eyes, the mouth, or both features were Thatcherised) on a set of event-related potential (ERP) components. Effects of inversion were found at the N170, P2 and P3b. Effects of eye condition were restricted to the N170 generated in the right hemisphere. Critically, an interaction of orientation and eye Thatcherisation was found for the P3b amplitude. Results from an individual with acquired prosopagnosia who can discriminate Thatcherised from typical faces but cannot categorise them or perceive the illusion (Mestry, Donnelly et al. in Neuropsychologia, 50, 3410-3418, 2012) only differed from typical participants at the P3b component. Findings suggest the P3b links most directly to the experience of the illusion. Overall, the study showed evidence consistent with both perceptual and decisional sources and the need to consider both in relation to configurality.

The late positive potential: a neural marker of the regulation of emotion-based approach-avoidance actions?

The ability to regulate our emotional responses is crucial to effective functioning in daily life. Whilst there has been extensive study of the brain potentials related to valenced stimuli, the neural basis of the ability to regulate actions elicited by these remains to be clarified. To address this, 40 volunteers undertook an approach-avoidance paradigm. In the congruent condition, participants approached pleasant and avoided unpleasant stimuli. In the incongruent condition, the opposite was the case, requiring the regulation of natural emotional response tendencies. Both behavioural and electrophysiological indices of emotional regulation were recorded. Congruency effects were observed at both the behavioural and electrophysiological level. Reaction times were faster and the LPP larger, when performing emotionally congruous relative to incongruous actions. Moreover, neural and behavioural effects were correlated. The current results suggest that the LPP congruency effect can be considered a neural marker of individual differences in emotion-driven action tendencies. We discuss whether this reflects emotion regulation, effort allocation, or correct mapping of stimulus response tendencies.

Short-term training in the Go/Nogo task: behavioural and neural changes depend on task demands.

Neural activity underlying executive functions is subject to modulation as a result of increasing cognitive demands and practice. In the present study, we examined these modulatory effects by varying task difficulty, as manipulated by reaction time deadline (RTD), on inhibitory control during a single Go/Nogo training session (8 blocks; 70% Go). Sixty adults were randomly assigned to one of three task difficulty conditions: High (n=20), Medium (n=20) and Low (n=20), with RTDs of 300, 500 or 1000 ms, respectively. Task performance, Event-related potentials (ERPs) and task-related arousal (indexed by skin conductance level) were examined for training effects. Results indicated that improvements in behavioural Go/Nogo proficiency were optimised during conditions of moderate rather than low or high inhibitory demands. An across-session increase in task-related arousal did not differ between conditions, indicating a generalised increase in the mobilisation of mental resources with time-on-task. In contrast, training-related changes in ERPs were dependent on task demands such that the Low task difficulty condition showed an enhanced centroparietal Nogo P2, while a training-induced augmentation in the Nogo>Go P3 effect was greater in the High than Medium condition. The High condition also showed the greatest reduction in the Nogo N1. Although further research is needed in this area, these findings implicate the potential key role of task difficulty in training inhibitory control and suggest that practice-related changes are reflected by qualitative changes in brain activity.

Varying task difficulty in the Go/Nogo task: the effects of inhibitory control, arousal, and perceived effort on ERP components.

Similar to other executive functions, inhibitory control is thought to be a dynamic process that can be influenced by variations in task difficulty. However, little is known about how different task parameters alter inhibitory performance and processing as a task becomes more difficult. The aim of this study was to investigate the influence of varying task difficulty, via manipulation of reaction time deadline (RTD), on measures of inhibitory control, perceived effort, and task-related arousal (indexed by skin conductance level). Sixty adults completed a visual Go/Nogo task (70% Go) after being randomly assigned to one of three task difficulty conditions: High, Medium and Low, with RTDs of 300, 500 or 1000 ms, respectively. Results revealed incremental increases in Go/Nogo errors and greater perceived effort with increasing difficulty. No condition differences were found for arousal, but the amplitude of the Nogo N2 increased and peaked earlier with increasing task difficulty. In contrast, the Nogo P3 effect was reduced in the High condition compared to the Low and Medium conditions. Finally, the amplitude of N1 and P2 showed differential effects, with Nogo N1 increasing with task difficulty, while the Nogo P2 decreased. This study provides valuable baseline behavioural and ERP data for appropriately manipulating difficulty (via RTD) in Go/Nogo tasks - highlighting the potential key role of not only the N2 and P3, but also the N1 and P2 components for task performance.

"Can waiting awaken the resting brain?" A comparison of waiting- and cognitive task-induced attenuation of very low frequency neural oscillations.

The default mode network (DMN) is characterised by coherent very low frequency (VLF) neural oscillations in the resting brain. The attenuation of this activity has been demonstrated following the transition from rest to performance of a broad range of cognitive goal-directed tasks. Whether the activity of resting state VLF oscillations is attenuated during non-cognitive goal-directed tasks such as waiting for rewarding outcomes is not known. This study examined the VLF EEG power from resting to performance of attention demanding task and two types of goal-directed waiting tasks. The association between the attenuation of VLF EEG power and Attention-Deficit/Hyperactivity Disorder (ADHD) symptoms was examined. Direct current EEG (DC-EEG) data were collected from 32 healthy young adults (half high and half low ADHD symptom scorers) during (i) a rest state, (ii) while performing a cognitive demanding reaction time task (2CRT), and (iii) while undertaking each of two different goal-directed waiting conditions: "forced-to-wait (FW)" and "choose-to-wait (CW)" tasks. The spatial distribution of VLF EEG power across scalp was similar to that seen in previous resting VLF EEG studies. Significant rest-to-task attenuation of VLF EEG power occurred during the 2CRT and the CW task, but not during the FW task. The association between self-ratings of ADHD symptoms and waiting-induced attenuation was not significant. This study suggests VLF EEG power attenuation that occurs following rest-to-task transition is not simply determined by changes in cognitive load. The goal-directed nature of a task, its motivated nature and/or the involvement of effortful attention may also contribute. Future studies should explore the attenuation of resting state VLF oscillations during waiting and impulsive choice.

Arousal-state modulation in children with AD/HD.

OBJECTIVE: To investigate the effect of arousal-state modulation, via manipulation of stimulus event-rate, on response inhibition in children with Attention-Deficit/Hyperactivity disorder (AD/HD) using behavioural and ERP measures. METHODS: Eighteen children with AD/HD, aged 7-14years, and 18 age-and sex-matched controls performed a cued visual Go/Nogo task (70% Go) with stimuli presented at fast, medium and slow event-rates. Task performance and ERPs to Warning, Go and Nogo stimuli, as well as preparation between the S1-S2 interval, were examined for group differences. RESULTS: AD/HD subjects displayed poorer response inhibition during the fast condition, accompanied by a reduced Nogo P3. Group differences during the fast rate extended to Warning cues, with the AD/HD group showing ERP evidence of atypical orienting/preparation, as indexed by the early and late CNV, and early sensory/attentive processing prior to S2. CONCLUSIONS: Although deficient response inhibition has been proposed as the core deficit in AD/HD, the results of the present study highlight the key role of energetic factors. Furthermore, group differences found to cues suggest that this effect extends to the processing of task-irrelevant stimuli. SIGNIFICANCE: This was the first ERP Go/Nogo task investigation using three event-rates, and the results support the theory that state factors may contribute to response inhibition deficits in AD/HD.