Neurophysiological evidence of motor preparation in inner speech and the effect of content predictability.

Self-generated overt actions are preceded by a slow negativity as measured by electroencephalogram, which has been associated with motor preparation. Recent studies have shown that this neural activity is modulated by the predictability of action outcomes. It is unclear whether inner speech is also preceded by a motor-related negativity and influenced by the same factor. In three experiments, we compared the contingent negative variation elicited in a cue paradigm in an active vs. passive condition. In Experiment 1, participants produced an inner phoneme, at which an audible phoneme whose identity was unpredictable was concurrently presented. We found that while passive listening elicited a late contingent negative variation, inner speech production generated a more negative late contingent negative variation. In Experiment 2, the same pattern of results was found when participants were instead asked to overtly vocalize the phoneme. In Experiment 3, the identity of the audible phoneme was made predictable by establishing probabilistic expectations. We observed a smaller late contingent negative variation in the inner speech condition when the identity of the audible phoneme was predictable, but not in the passive condition. These findings suggest that inner speech is associated with motor preparatory activity that may also represent the predicted action-effects of covert actions.

Estimating statistical power for ERP studies using the auditory N1, Tb, and P2 components.

The N1, Tb, and P2 components of the event-related potential (ERP) are thought to reflect the sequential processing of auditory stimuli in the human brain. Despite their extensive use in biological, cognitive, and clinical neuroscience, there are no guidelines for how to appropriately power ERP studies using these components. In the present study, we investigated how the number of trials, number of participants, effect magnitude, and study design influenced statistical power. Using Monte Carlo simulations of ERP data from a passive listening task, we determined the probability of finding a statistically significant effect in 58,900 experiments repeated 1,000 times each. We found that as the number of trials, number of participants, and effect magnitude increased, so did statistical power. We also found that increasing the number of trials had a bigger effect on statistical power for within-subject designs than for between-subject designs, and that within-subject designs required a smaller number of trials and participants to provide the same level of statistical power for a given effect magnitude than between-subject designs. These results show that it is important to carefully consider these factors when designing ERP studies, rather than relying on tradition or anecdotal evidence. To improve the robustness and reproducibility of ERP research, we have built an online statistical power calculator (https://bradleynjack.shinyapps.io/ErpPowerCalculator), which we hope will allow researchers to estimate the statistical power of previous studies, as well as help them design appropriately-powered studies in the future.

Disrupted auditory N1, theta power and coherence suppression to willed speech in people with schizophrenia.

The phenomenon of sensory self-suppression - also known as sensory attenuation - occurs when a person generates a perceptible stimulus (such as a sound) by performing an action (such as speaking). The sensorimotor control system is thought to actively predict and then suppress the vocal sound in the course of speaking, resulting in lowered cortical responsiveness when speaking than when passively listening to an identical sound. It has been hypothesized that auditory hallucinations in schizophrenia result from a reduction in self-suppression due to a disruption of predictive mechanisms required to anticipate and suppress a specific, self-generated sound. It has further been hypothesized that this suppression is evident primarily in theta band activity. Fifty-one people, half of whom had a diagnosis of schizophrenia, were asked to repeatedly utter a single syllable, which was played back to them concurrently over headphones while EEG was continuously recorded. In other conditions, recordings of the same spoken syllables were played back to participants while they passively listened, or were played back with their onsets preceded by a visual cue. All participants experienced these conditions with their voice artificially shifted in pitch and also with their unaltered voice. Suppression was measured using event-related potentials (N1 component), theta phase coherence and power. We found that suppression was generally reduced on all metrics in the patient sample, and when voice alteration was applied. We additionally observed reduced theta coherence and power in the patient sample across all conditions. Visual cueing affected theta coherence only. In aggregate, the results suggest that sensory self-suppression of theta power and coherence is disrupted in schizophrenia.

The Role of Action-Effect Contingency on Sensory Attenuation in the Absence of Movement.

Stimuli that have been generated by a person's own willed motor actions generally elicit a suppressed electrophysiological, as well as phenomenological, response compared with identical stimuli that have been externally generated. This well-studied phenomenon, known as sensory attenuation, has mostly been studied by comparing ERPs evoked by self-initiated and externally generated sounds. However, most studies have assumed a uniform action-effect contingency, in which a motor action leads to a resulting sensation 100% of the time. In this study, we investigated the effect of manipulating the probability of action-effect contingencies on the sensory attenuation effect. In Experiment 1, participants watched a moving, marked tickertape while EEG was recorded. In the full-contingency (FC) condition, participants chose whether to press a button by a certain mark on the tickertape. If a button press had not occurred by the mark, a sound would be played a second later 100% of the time. If the button was pressed before the mark, the sound was not played. In the no-contingency (NC) condition, participants observed the same tickertape; in contrast, however, if participants did not press the button by the mark, a sound would occur only 50% of the time (NC-inaction). Furthermore, in the NC condition, if a participant pressed the button before the mark, a sound would also play 50% of the time (NC-action). In Experiment 2, the design was identical, except that a willed action (as opposed to a willed inaction) triggered the sound in the FC condition. The results were consistent across the two experiments: Although there were no differences in N1 amplitude between conditions, the amplitude of the Tb and P2 components were smaller in the FC condition compared with the NC-inaction condition, and the amplitude of the P2 component was also smaller in the FC condition compared with the NC-action condition. The results suggest that the effect of contingency on electrophysiological indices of sensory attenuation may be indexed primarily by the Tb and P2 components, rather than the N1 component which is most commonly studied.

The Relationship Between Affective Visual Mismatch Negativity and Interpersonal Difficulties Across Autism and Schizotypal Traits.

Sensory deficits are a feature of autism and schizophrenia, as well as the upper end of their non-clinical spectra. The mismatch negativity (MMN), an index of pre-attentive auditory processing, is particularly sensitive in detecting such deficits; however, little is known about the relationship between the visual MMN (vMMN) to facial emotions and autism and schizophrenia spectrum symptom domains. We probed the vMMN to happy, sad, and neutral faces in 61 healthy adults (18-40 years, 32 female), and evaluated their degree of autism and schizophrenia spectrum traits using the Autism Spectrum Quotient (AQ) and Schizotypal Personality Questionnaire (SPQ). The vMMN to happy faces was significantly larger than the vMMNs to sad and neutral faces. The vMMN to happy faces was associated with interpersonal difficulties as indexed by AQ Communication and Attention to Detail subscales, and SPQ associated with more interpersonal difficulties. These data suggest that pre-attentive processing of positive affect might be more specific to the interpersonal features associated with autism and schizophrenia. These findings add valuable insights into the growing body of literature investigating symptom-specific neurobiological markers of autism and schizophrenia spectrum conditions.

Movement Planning Determines Sensory Suppression: An Event-related Potential Study.

Sensory suppression refers to the phenomenon that sensory input generated by our own actions, such as moving a finger to press a button to hear a tone, elicits smaller neural responses than sensory input generated by external agents. This observation is usually explained via the internal forward model in which an efference copy of the motor command is used to compute a corollary discharge, which acts to suppress sensory input. However, because moving a finger to press a button is accompanied by neural processes involved in preparing and performing the action, it is unclear whether sensory suppression is the result of movement planning, movement execution, or both. To investigate this, in two experiments, we compared ERPs to self-generated tones that were produced by voluntary, semivoluntary, or involuntary button-presses, with externally generated tones that were produced by a computer. In Experiment 1, the semivoluntary and involuntary button-presses were initiated by the participant or experimenter, respectively, by electrically stimulating the median nerve in the participant's forearm, and in Experiment 2, by applying manual force to the participant's finger. We found that tones produced by voluntary button-presses elicited a smaller N1 component of the ERP than externally generated tones. This is known as N1-suppression. However, tones produced by semivoluntary and involuntary button-presses did not yield significant N1-suppression. We also found that the magnitude of N1-suppression linearly decreased across the voluntary, semivoluntary, and involuntary conditions. These results suggest that movement planning is a necessary condition for producing sensory suppression. We conclude that the most parsimonious account of sensory suppression is the internal forward model.

Sensory attenuation in the absence of movement: Differentiating motor action from sense of agency.

Sensory attenuation is the phenomenon that stimuli generated by willed motor actions elicit a smaller neurophysiological response than those generated by external sources. It has mostly been investigated in the auditory domain, by comparing ERPs evoked by self-initiated (active condition) and externally-generated (passive condition) sounds. The mechanistic basis of sensory attenuation has been argued to involve a duplicate of the motor command being used to predict sensory consequences of self-generated movements. An alternative possibility is that the effect is driven by between-condition differences in participants' sense of agency over the sound. In this paper, we disambiguated the effects of motor-action and sense of agency on sensory attenuation with a novel experimental paradigm. In Experiment 1, participants watched a moving, marked tickertape while EEG was recorded. In the active condition, participants chose whether to press a button by a certain mark on the tickertape. If a button-press had not occurred by the mark, then a tone would be played 1 s later. If the button was pressed prior to the mark, the tone was not played. In the passive condition, participants passively watched the animation, and were informed about whether a tone would be played on each trial. The design for Experiment 2 was identical, except that the contingencies were reversed (i.e., a button-press by the mark led to a tone). The results were consistent across the two experiments: while there were no differences in N1 amplitude between the active and passive conditions, the amplitude of the Tb component was suppressed in the active condition. The amplitude of the P2 component was enhanced in the active condition in both Experiments 1 and 2. These results suggest that motor-actions and sense of agency have differential effects on sensory attenuation to sounds and are indexed with different ERP components.

Sensory attenuation is modulated by the contrasting effects of predictability and control.

Self-generated stimuli have been found to elicit a reduced sensory response compared with externally-generated stimuli. However, much of the literature has not adequately controlled for differences in the temporal predictability and temporal control of stimuli. In two experiments, we compared the N1 (and P2) components of the auditory-evoked potential to self- and externally-generated tones that differed with respect to these two factors. In Experiment 1 (n = 42), we found that increasing temporal predictability reduced N1 amplitude in a manner that may often account for the observed reduction in sensory response to self-generated sounds. We also observed that reducing temporal control over the tones resulted in a reduction in N1 amplitude. The contrasting effects of temporal predictability and temporal control on N1 amplitude meant that sensory attenuation prevailed when controlling for each. Experiment 2 (n = 38) explored the potential effect of selective attention on the results of Experiment 1 by modifying task requirements such that similar levels of attention were allocated to the visual stimuli across conditions. The results of Experiment 2 replicated those of Experiment 1, and suggested that the observed effects of temporal control and sensory attenuation were not driven by differences in attention. Given that self- and externally-generated sensations commonly differ with respect to both temporal predictability and temporal control, findings of the present study may necessitate a re-evaluation of the experimental paradigms used to study sensory attenuation.

#EEGManyLabs: Investigating the replicability of influential EEG experiments.

There is growing awareness across the neuroscience community that the replicability of findings about the relationship between brain activity and cognitive phenomena can be improved by conducting studies with high statistical power that adhere to well-defined and standardised analysis pipelines. Inspired by recent efforts from the psychological sciences, and with the desire to examine some of the foundational findings using electroencephalography (EEG), we have launched #EEGManyLabs, a large-scale international collaborative replication effort. Since its discovery in the early 20th century, EEG has had a profound influence on our understanding of human cognition, but there is limited evidence on the replicability of some of the most highly cited discoveries. After a systematic search and selection process, we have identified 27 of the most influential and continually cited studies in the field. We plan to directly test the replicability of key findings from 20 of these studies in teams of at least three independent laboratories. The design and protocol of each replication effort will be submitted as a Registered Report and peer-reviewed prior to data collection. Prediction markets, open to all EEG researchers, will be used as a forecasting tool to examine which findings the community expects to replicate. This project will update our confidence in some of the most influential EEG findings and generate a large open access database that can be used to inform future research practices. Finally, through this international effort, we hope to create a cultural shift towards inclusive, high-powered multi-laboratory collaborations.

Distinguishing schizophrenia spectrum from non-spectrum disorders among young patients with first episode psychosis and at high clinical risk: The role of basic self-disturbance and neurocognition.

INTRODUCTION: The distinction between the schizophrenia spectrum and other types of disorders may be clinically relevant in terms of its predictive validity as suggested by studies showing schizophrenia spectrum patients have more unfavourable outcomes compared to other psychotic disorders. The present study aimed to investigate whether basic self-disturbances and neurocognitive processes that have been linked to psychosis risk have discriminative power for schizophrenia spectrum disorders in patients presenting with first episode psychosis (FEP) and at ultra-high risk for psychosis (UHR). METHODS: 38 FEP patients, 48 UHR patients, and 33 healthy controls were assessed for basic self-disturbances (using the Examination of Anomalous Self-Experience, EASE, interview), source monitoring and aberrant salience (behavioural tasks to measure neurocognitive constructs). Clinical groups were divided into patients with schizophrenia spectrum disorders and those with other non-spectrum disorders and were further compared on measures controlling for symptom severity and age. RESULTS: Basic self-disturbances distinguished schizophrenia spectrum from non-spectrum disorders in the 'FEP only' sample, F = 19.76, p < 0.001, η2partial = 0.37, and also in the combined UHR/FEP sample, F = 23.56, p < 0.001, η2partial = 0.22. Additionally, some processes related to source monitoring deficits were elevated in schizophrenia spectrum disorders. In contrast, the two groups (schizophrenia spectrum vs other diagnoses) performed similarly in aberrant salience tasks. Comparable results were obtained for analyses performed with an FEP/UHR combined sample and the 'FEP only' sample. DISCUSSION: Basic self-disturbances at the phenomenological level and source monitoring deficits on the neurocognitive level may be useful in identifying risk of schizophrenia spectrum disorders at the earliest clinical presentation.

Inner speech is accompanied by a temporally-precise and content-specific corollary discharge.

When we move our articulator organs to produce overt speech, the brain generates a corollary discharge that acts to suppress the neural and perceptual responses to our speech sounds. Recent research suggests that inner speech - the silent production of words in one's mind - is also accompanied by a corollary discharge. Here, we show that this corollary discharge contains information about the temporal and physical properties of inner speech. In two experiments, participants produced an inner phoneme at a precisely-defined moment in time. An audible phoneme was presented 300 ms before, concurrently with, or 300 ms after participants produced the inner phoneme. We found that producing the inner phoneme attenuated the N1 component of the event-related potential - an index of auditory cortex processing - but only when the inner and audible phonemes occurred concurrently and matched on content. If the audible phoneme was presented before or after the production of the inner phoneme, or if the inner phoneme did not match the content of the audible phoneme, there was no attenuation of the N1. These results suggest that inner speech is accompanied by a temporally-precise and content-specific corollary discharge. We conclude that these results support the notion of a functional equivalence between the neural processes that underlie the production of inner and overt speech, and may provide a platform for identifying inner speech abnormalities in disorders in which they have been putatively associated, such as schizophrenia.

Frontal slow wave resting EEG power is higher in individuals at Ultra High Risk for psychosis than in healthy controls but is not associated with negative symptoms or functioning.

Decreased brain activity in the frontal region, as indicated by increased slow wave EEG power measured by electrodes place on the skull over this area, in association with negative symptoms has previously been shown to distinguish ultra-high risk (UHR) individuals who later transitioned to psychosis (UHR-P) from those who did not transition (UHR-NP). The aims of the current study were to: 1) replicate these results and 2) investigate whether similar association between increased frontal slow wave activity and functioning shows any value in the prediction of transition to psychosis in UHR individuals. The brain activity, recorded using EEG, of 44 UHR individuals and 38 healthy controls was included in the analyses. Symptom severity was assessed in UHR participants and functioning was measured in both groups. The power in the theta frequency band in the frontal region of UHR individuals was higher than in controls. However, there was no difference between the UHR-P and the UHR-NP groups, and no change in slow frequency power following transition to psychosis. The correlation between delta frequency power and negative symptoms previously observed was not present in our UHR cohort, and there was no association between frontal delta or theta and functioning in either group. Increased delta power was rather correlated with depressive symptoms in the UHR group. Future research will be needed to better understand when, in the course of the illness, does the slow wave activity in the frontal area becomes impaired.

Semantic prediction-errors are context-dependent: An ERP study.

The human brain is an efficient, adaptive, and predictive machine, constructing a generative model of the environment that we then perceive and become conscious of. Here, we show that different types of prediction-errors - the discrepancies between top-down expectations and bottom-up sensory input - are integrated across processing levels and sensory modalities of the cortical hierarchy. We designed a novel, hybrid protocol in which five prediction-establishing sounds were played in rapid succession (e.g., "meow", "meow", "meow", etc.), followed by either a standard (e.g., "meow") or a deviant (e.g., "woof") prime sound, then a visual target word that was either congruent or incongruent (e.g., "cat" or "dog") with the prime sound. We found that the deviants elicited a more negative voltage than the standards at about 150 ms - the mismatch negativity (MMN), an event-related potential (ERP) sensitive to low-level perceptual violations - and that the incongruent words elicited a more negative voltage than the congruent words at about 350 ms - the N400, an ERP sensitive to high-level semantic violations. We also found that the N400 was context-dependent: the N400 was larger when the target words were preceded by a standard than a deviant. Our results suggest that perceptual prediction-errors modulate subsequent semantic prediction-errors. We conclude that our results are consistent with one of the most important assumptions of predictive coding theories: hierarchical prediction-error processing.

Impaired mismatch negativity to frequency deviants in individuals at ultra-high risk for psychosis, and preliminary evidence for further impairment with transition to psychosis.

BACKGROUND: There is evidence to suggest that people with established psychotic disorders show impairments in the mismatch negativity induced by a frequency-deviant sound (fMMN), and that these impairments worsen with the deterioration of psychotic symptoms. This study aimed to test whether individuals at ultra-high risk (UHR) for psychosis show pre-morbid impairments in fMMN, and if so, whether fMMN continues to deteriorate with transition to psychosis. METHOD: fMMN was recorded in a cohort of UHR individuals (n=42) and compared to healthy controls (n=29). Of the 27 UHR participants who returned for a second EEG session, six participants had transitioned to psychosis by 12-month follow-up (UHR-T) and were compared to the 21 participants who did not transition (UHR-NT). RESULTS: fMMN amplitude was significantly reduced, relative to healthy controls, in the UHR cohort. Furthermore, UHR-T individuals showed a significant decrease in fMMN amplitude over the period from baseline to post-transition; this reduction was not observed in UHR-NT. CONCLUSIONS: These results suggest that fMMN is abnormal in UHR individuals, as has repeatedly been found previously in people with established psychotic disorders. The finding that fMMN impairment worsens with transition to psychosis is consistent with the staging model of psychosis; however, caution must be taken in interpreting these findings, given the extremely small sample size of the UHR-T group.

Neurophysiological evidence of efference copies to inner speech.

Efference copies refer to internal duplicates of movement-producing neural signals. Their primary function is to predict, and often suppress, the sensory consequences of willed movements. Efference copies have been almost exclusively investigated in the context of overt movements. The current electrophysiological study employed a novel design to show that inner speech - the silent production of words in one's mind - is also associated with an efference copy. Participants produced an inner phoneme at a precisely specified time, at which an audible phoneme was concurrently presented. The production of the inner phoneme resulted in electrophysiological suppression, but only if the content of the inner phoneme matched the content of the audible phoneme. These results demonstrate that inner speech - a purely mental action - is associated with an efference copy with detailed auditory properties. These findings suggest that inner speech may ultimately reflect a special type of overt speech.

Do early neural correlates of visual consciousness show the oblique effect? A binocular rivalry and event-related potential study.

When dissimilar images are presented one to each eye, we do not see both images; rather, we see one at a time, alternating unpredictably. This is called binocular rivalry, and it has recently been used to study brain processes that correlate with visual consciousness, because perception changes without any change in the sensory input. Such studies have used various types of images, but the most popular have been gratings: sets of bright and dark lines of orthogonal orientations presented one to each eye. We studied whether using cardinal rival gratings (vertical, 0°, and horizontal, 90°) versus oblique rival gratings (left-oblique, -45°, and right-oblique, 45°) influences early neural correlates of visual consciousness, because of the oblique effect: the tendency for visual performance to be greater for cardinal gratings than for oblique gratings. Participants viewed rival gratings and pressed keys indicating which of the two gratings they perceived, was dominant. Next, we changed one of the gratings to match the grating shown to the other eye, yielding binocular fusion. Participants perceived the rivalry-to-fusion change to the dominant grating and not to the other, suppressed grating. Using event-related potentials (ERPs), we found neural correlates of visual consciousness at the P1 for both sets of gratings, as well as at the P1-N1 for oblique gratings, and we found a neural correlate of the oblique effect at the N1, but only for perceived changes. These results show that the P1 is the earliest neural activity associated with visual consciousness and that visual consciousness might be necessary to elicit the oblique effect.

Brain activity from stimuli that are not perceived: Visual mismatch negativity during binocular rivalry suppression.

Predictive coding explains visual perception as the result of an interaction between bottom-up sensory input and top-down generative models at each level of the visual hierarchy. Evidence for this comes from the visual mismatch negativity (vMMN): a more negative ERP for rare, unpredictable visual stimuli-deviants, than for frequent, predictable visual stimuli-standards. Here, we show that the vMMN does not require conscious experience. We measured the vMMN from monocular luminance-decrement deviants that were either perceived or not during binocular rivalry dominance or suppression, respectively. We found that both sorts of deviants elicited the vMMN at about 250 ms after stimulus onset, with perceived deviants eliciting a bigger vMMN than not-perceived deviants. These results show that vMMN occurs in the absence of consciousness, and that consciousness enhances the processing underlying vMMN. We conclude that generative models of visual perception are tested, even when sensory input for those models is not perceived.

Goal-Directed and Habit-Like Modulations of Stimulus Processing during Reinforcement Learning.

Recent research has shown that perceptual processing of stimuli previously associated with high-value rewards is automatically prioritized even when rewards are no longer available. It has been hypothesized that such reward-related modulation of stimulus salience is conceptually similar to an "attentional habit." Recording event-related potentials in humans during a reinforcement learning task, we show strong evidence in favor of this hypothesis. Resistance to outcome devaluation (the defining feature of a habit) was shown by the stimulus-locked P1 component, reflecting activity in the extrastriate visual cortex. Analysis at longer latencies revealed a positive component (corresponding to the P3b, from 550-700 ms) sensitive to outcome devaluation. Therefore, distinct spatiotemporal patterns of brain activity were observed corresponding to habitual and goal-directed processes. These results demonstrate that reinforcement learning engages both attentional habits and goal-directed processes in parallel. Consequences for brain and computational models of reinforcement learning are discussed.SIGNIFICANCE STATEMENT The human attentional network adapts to detect stimuli that predict important rewards. A recent hypothesis suggests that the visual cortex automatically prioritizes reward-related stimuli, driven by cached representations of reward value; that is, stimulus-response habits. Alternatively, the neural system may track the current value of the predicted outcome. Our results demonstrate for the first time that visual cortex activity is increased for reward-related stimuli even when the rewarding event is temporarily devalued. In contrast, longer-latency brain activity was specifically sensitive to transient changes in reward value. Therefore, we show that both habit-like attention and goal-directed processes occur in the same learning episode at different latencies. This result has important consequences for computational models of reinforcement learning.

Cross-modal symbolic processing can elicit either an N2 or a protracted N2/N400 response.

A cross-modal symbolic paradigm was used to elicit EEG activity related to semantic incongruence. Twenty-five undergraduate students viewed pairings of visual lexical cues (e.g., DOG) with congruent (50% of trials) or incongruent (50%) auditory nonlexical stimuli (animal vocalizations; e.g., sound of a dog woofing or a cat meowing). In one condition, many different pairs of congruent/incongruent stimuli were shown, whereas in a second condition only two pairs of stimuli were repeatedly shown. A typical N400-like pattern of incongruence-related activity (including activity in the N2 time window) was evident in the condition using many stimuli, whereas the incongruence-related activity in the two-stimuli condition was confined to differential N2-like activity. A supplementary analysis excluded stimulus characteristics as the source of this differential activity between conditions. We found that a single individual performing a fixed task can demonstrate either a protracted N400-like pattern of activity or a more temporally focused N2-like pattern of activity in response to the same stimulus, which suggests that the N2 may be a precursor to the protracted N400 response.