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.

How top-down and bottom-up attention modulate risky choice.

We examine how bottom-up (or stimulus-driven) and top-down (or goal-driven) processes govern the distribution of attention in risky choice. In three experiments, participants chose between a certain payoff and the chance of receiving a payoff drawn randomly from an array of eight numbers. We tested the hypothesis that initial attention is driven by perceptual properties of the stimulus (e.g., font size of the numbers), but subsequent choice is goal-driven (e.g., win the best outcome). Two experiments in which task framing (goal driven) and font size (stimulus driven) were manipulated demonstrated that payoffs with the highest values and the largest font sizes had the greatest impact on choice. The third experiment added a number in large font to the array, which could not be an outcome of the gamble (i.e., a distractor). Eye movement and choice data indicated that although the distractor attracted attention, it had no influence on option selection. Together with computational modeling analyses, the results suggest that perceptual salience can induce bottom-up effects of overt selection but that the perceived value of information is the crucial arbiter of intentional control over risky choice.

The Neural Bases of Action-Outcome Learning in Humans.

From an associative perspective the acquisition of new goal-directed actions requires the encoding of specific action-outcome (AO) associations and, therefore, sensitivity to the validity of an action as a predictor of a specific outcome relative to other events. Although competitive architectures have been proposed within associative learning theory to achieve this kind of identity-based selection, whether and how these architectures are implemented by the brain is still a matter of conjecture. To investigate this issue, we trained human participants to encode various AO associations while undergoing functional neuroimaging (fMRI). We then degraded one AO contingency by increasing the probability of the outcome in the absence of its associated action while keeping other AO contingencies intact. We found that this treatment selectively reduced performance of the degraded action. Furthermore, when a signal predicted the unpaired outcome, performance of the action was restored, suggesting that the degradation effect reflects competition between the action and the context for prediction of the specific outcome. We used a Kalman filter to model the contribution of different causal variables to AO learning and found that activity in the medial prefrontal cortex (mPFC) and the dorsal anterior cingulate cortex (dACC) tracked changes in the association of the action and context, respectively, with regard to the specific outcome. Furthermore, we found the mPFC participated in a network with the striatum and posterior parietal cortex to segregate the influence of the various competing predictors to establish specific AO associations.SIGNIFICANCE STATEMENT Humans and other animals learn the consequences of their actions, allowing them to control their environment in a goal-directed manner. Nevertheless, it is unknown how we parse environmental causes from the effects of our own actions to establish these specific action-outcome (AO) relationships. Here, we show that the brain learns the causal structure of the environment by segregating the unique influence of actions from other causes in the medial prefrontal and anterior cingulate cortices and, through a network of structures, including the caudate nucleus and posterior parietal cortex, establishes the distinct causal relationships from which specific AO associations are formed.

Attentional capture by signals of reward persists following outcome devaluation.

Attention, the mechanism that prioritizes stimuli in the environment for further processing, plays an important role in behavioral choice. In the present study, we investigated the automatic orienting of attention to cues that signal reward. Such attentional capture occurs despite negative consequences, and we investigated whether this counterproductive and reflexive behavior would persist following outcome devaluation. Thirsty participants completed a visual search task in which the color of a distractor stimulus in the search display signaled whether participants would earn water or potato chips for making a rapid eye movement to a diamond target, but looking at the colored distractor was punished by omission of the signaled reward. Nevertheless, participants looked at the water-signaling distractor more frequently than the chip-signaling distractor. Half the participants then drank water ad libitum before continuing with the visual search task. Although the water was now significantly less desirable for half of the participants, there was no difference between groups in the tendency for the water-signaling distractor to capture attention. These findings suggest that once established, counterproductive attentional bias to signals of reward persists even when those outcomes are no longer valuable. This suggests a "habit-like" attentional mechanism that prioritizes reward stimuli in the environment for further action, regardless of whether those stimuli are aligned with current goals or currently desired.

Learning to like triangles: A longitudinal investigation of evaluative conditioning in infancy.

There is tentative evidence that infants can learn preferences through evaluative conditioning to socioemotional stimuli. However, the early development of evaluative conditioning and the factors that may explain infants' capacity to learn through evaluative conditioning are not well understood. Infants (N = 319; 50.2% boys) participated in a longitudinal study where an evaluative conditioning paradigm using socioemotional stimuli was conducted on two occasions (when infants were 7 and 14 months old, on average). We tested whether repeatedly pairing neutral stimuli (triangular and square shapes) with affective stimuli (angry and happy faces) affects infants' preferences for these shapes. At both timepoints, the majority of infants did not choose the shape that was paired with happy faces, indicating that, in general, learning through evaluative conditioning was not present. However, as expected, individual differences were evident such that infants who spent more time fixating on faces compared to shapes (face-preferrers) during the conditioning trials were significantly more likely than non-face-preferrers to choose the shape paired with happy faces, and this effect strengthened with increasing age.

Still connecting the dots: An investigation into infants' attentional bias to threat using an eye-tracking task.

An attentional bias toward threat has been theorized to be a normative aspect of infants' threat and safety learning, and an indicator of risk for internalizing psychopathology in older populations. To date, only four studies have examined this bias using the dot-probe task in infancy and the findings are mixed. We extended the literature by examining patterns of attention to threat in a culturally and linguistically diverse sample of infants aged 5-11 months old (N = 151) using all measures previously employed in the infant dot-probe literature. Given that an attentional bias toward threat is associated with higher risk of developing anxiety disorders later in life, we also examined how negative affect-an early correlate of later anxiety disorders-is related to attentional bias toward threat in infancy. This study was the first to use a consistent measure of negative affect across the whole sample. An eye-tracking dot-probe task was used to examine attentional bias toward threat (i.e., angry faces) relative to positive (i.e., happy faces) stimuli. Results showed that an attention bias to threat was not characteristic of infants at this age, and negative affect did not moderate the putative relationship between attention and emotional faces (angry, happy). These findings therefore suggest that attention biases to socio-emotional threat may not have emerged by 11 months old.

Reward Rapidly Enhances Visual Perception.

Rewards exert a deep influence on our cognition and behavior. Here, we used a paradigm in which reward information was provided at either encoding or retrieval of a brief, masked stimulus to show that reward can also rapidly modulate perceptual encoding of visual information. Experiment 1 (n = 30 adults) showed that participants' response accuracy was enhanced when a to-be-encoded grating signaled high reward relative to low reward, but only when the grating was presented very briefly and participants reported that they were not consciously aware of it. Experiment 2 (n = 29 adults) showed that there was no difference in participants' response accuracy when reward information was instead provided at the stage of retrieval, ruling out an explanation of the reward-modulation effect in terms of differences in motivated retrieval. Taken together, our findings provide behavioral evidence consistent with a rapid reward modulation of visual perception, which may not require consciousness.

Reward-Related Attentional Capture Moderates the Association between Fear-Driven Motives and Heavy Drinking.

BACKGROUND: To date, there has been little investigation on how motivational and cognitive mechanisms interact to influence problematic drinking behaviours. Towards this aim, the current study examined whether reward-related attentional capture is associated with reward, fear (relief), and habit drinking motives, and further, whether it interacts with these motives in relation to problematic drinking patterns. METHODS: Ninety participants (mean age = 34.8 years, SD = 9.1, 54% male) who reported having consumed alcohol in the past month completed an online visual search task that measured reward-related attentional capture as well as the Habit Reward Fear Scale, a measure of drinking motives. Participants also completed measures of psychological distress, impulsivity, compulsive drinking, and consumption items of Alcohol Use Disorders Identification Test. Regression analyses examined the associations between motives for alcohol consumption and reward-related attentional capture, as well as the associations between reward-related attentional capture, motives, and their interaction, with alcohol consumption and problems. RESULTS: Greater reward-related attentional capture was associated with greater reward motives. Further, reward-related attentional capture also interacted with fear motives in relation to alcohol consumption. Follow-up analyses showed that this interaction was driven by greater fear motives being associated with heavier drinking among those with lower reward-related attentional capture (i.e., "goal-trackers"). CONCLUSION: These findings have implications for understanding how cognition may interact with motives in association with problematic drinking. Specifically, the findings highlight different potential pathways to problematic drinking according to an individual's cognitive-motivational profile and may inform tailored interventions to target profile-specific mechanisms. Finally, these findings offer support for contemporary models of addiction that view excessive goal-directed behaviour under negative affect as a critical contributor to addictive behaviours.

Eating restraint is associated with reduced attentional capture by signals of valuable food reward.

Previous paradigms used to examine attentional distraction by task-irrelevant food words and food images were not suited for the investigation of involuntary and automatic attentional capture. In the current experiments we adapted a well-established visual-search paradigm (with eye tracking) to investigate involuntary attentional capture by food and drink rewards. We first used a satiety procedure to manipulate relative preference for different food and drink outcomes (potato chips and water in Experiment 1 and popcorn and chocolate Smarties in Experiment 2). Participants then performed the visual-search task where a coloured distractor signalled on each trial which of the two food and drink rewards was available for successful identification of the target. The signalled reward was cancelled, however, if any eye gaze was registered on the distractor. Participants were therefore motivated to try and control the automatic orienting of attention towards cues signalling valuable outcomes, in order to earn those outcomes. In both experiments we found that attention was more often captured by the distractor signalling the valuable (non-sated) outcome, replicating previous studies using this paradigm with monetary rewards. We also found that those scoring high on eating restraint (as measured with the Dutch Eating Behavior Questionnaire) were better at controlling reflexive orienting of attention to desirable food rewards. This paradigm offers a novel approach for understanding how reflexive attention and control relate to conflicts in everyday life around distracting food cues, and the moderating role of dietary restraint.

Physiological and subjective validation of a novel stress procedure: The Simple Singing Stress Procedure.

Laboratory stress-induction procedures have been critical in illuminating the effects of stress on human health, cognition, and functioning. Here, we present a novel stress induction procedure, the Simple Singing Stress Procedure (SSSP), that overcomes some of the practical challenges and conceptual limitations of existing procedures in measuring the causal influence of stress on psychological variables. In the stress condition of the SSSP, participants were instructed to sing a song in front of the experimenter while being video- and audio-recorded. Participants were also informed that they would have to sing again at the end of the experiment, and that this second performance would later be assessed by a panel of experimenters. Participants in a no-stress condition instead read lyrics in each phase. Our findings revealed that participants in the stress condition showed significantly higher blood pressure immediately following the initial singing session, as well as heightened salivary cortisol at a latency consistent with the initial singing session, than those in the no-stress condition. Our stress procedure also generated elevations in self-reported stress ratings immediately after the first singing session and subsequently in anticipation of the second singing session, relative to the no-stress condition. Collectively, these findings suggest that the SSSP is a simple and effective stress induction procedure that may be a promising alternative to existing protocols.

Compulsivity is measurable across distinct psychiatric symptom domains and is associated with familial risk and reward-related attentional capture.

BACKGROUND: Compulsivity can be seen across various mental health conditions and refers to a tendency toward repetitive habitual acts that are persistent and functionally impairing. Compulsivity involves dysfunctional reward-related circuitry and is thought to be significantly heritable. Despite this, its measurement from a transdiagnostic perspective has received only scant research attention. Here we examine both the psychometric properties of a recently developed compulsivity scale, as well as its relationship with compulsive symptoms, familial risk, and reward-related attentional capture. METHODS: Two-hundred and sixty individuals participated in the study (mean age = 36.0 [SD = 10.8] years; 60.0% male) and completed the Cambridge-Chicago Compulsivity Trait Scale (CHI-T), along with measures of psychiatric symptoms and family history thereof. Participants also completed a task designed to measure reward-related attentional capture (n = 177). RESULTS: CHI-T total scores had a normal distribution and acceptable Cronbach's alpha (0.84). CHI-T total scores correlated significantly and positively (all p < 0.05, Bonferroni corrected) with Problematic Usage of the Internet, disordered gambling, obsessive-compulsive symptoms, alcohol misuse, and disordered eating. The scale was correlated significantly with history of addiction and obsessive-compulsive related disorders in first-degree relatives of participants and greater reward-related attentional capture. CONCLUSIONS: These findings suggest that the CHI-T is suitable for use in online studies and constitutes a transdiagnostic marker for a range of compulsive symptoms, their familial loading, and related cognitive markers. Future work should more extensively investigate the scale in normative and clinical cohorts, and the role of value-modulated attentional capture across compulsive disorders.

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.

Capture and Control: Working Memory Modulates Attentional Capture by Reward-Related Stimuli.

Physically salient but task-irrelevant distractors can capture attention in visual search, but resource-dependent, executive-control processes can help reduce this distraction. However, it is not only physically salient stimuli that grab our attention: Recent research has shown that reward history also influences the likelihood that stimuli will capture attention. Here, we investigated whether resource-dependent control processes modulate the effect of reward on attentional capture, much as for the effect of physical salience. To this end, we used eye tracking with a rewarded visual search task and compared performance under conditions of high and low working memory load. In two experiments, we demonstrated that oculomotor capture by high-reward distractor stimuli is enhanced under high memory load. These results highlight the role of executive-control processes in modulating distraction by reward-related stimuli. Our findings have implications for understanding the neurocognitive processes involved in real-life conditions in which reward-related stimuli may influence behavior, such as addiction.

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.

Act Now, Play Later: Temporal Expectations Regarding the Onset of Self-initiated Sensations Can Be Modified with Behavioral Training.

Mechanisms of motor-sensory prediction are dependent on expectations regarding when self-generated feedback will occur. Existing behavioral and electrophysiological research suggests that we have a default expectation for immediate sensory feedback after executing an action. However, studies investigating the adaptability of this temporal expectation have been limited in their ability to differentiate modified expectations per se from effects of stimulus repetition. Here, we use a novel, within-participant procedure that allowed us to disentangle the effect of repetition from expectation and allowed us to determine whether the default assumption for immediate feedback is fixed and resistant to modification or is amenable to change with experience. While EEG was recorded, 45 participants completed a task in which they repeatedly pressed a button to produce a tone that occurred immediately after the button press (immediate training) or after a 100-msec delay (delayed training). The results revealed significant differences in the patterns of cortical change across the two training conditions. Specifically, there was a significant reduction in the cortical response to tones across delayed training blocks but no significant change across immediate training blocks. Furthermore, experience with delayed training did not result in increased cortical activity in response to immediate feedback. These findings suggest that experience with action-sensation delays broadens the window of temporal expectations, allowing for the simultaneous anticipation of both delayed and immediate motor-sensory feedback. This research provides insights into the mechanisms underlying motor-sensory prediction and may represent a novel therapeutic avenue for psychotic symptoms, which are ostensibly associated with sensory prediction abnormalities.

Anterior Temporal Lobe Tracks the Formation of Prejudice.

Despite advances in understanding the brain structures involved in the expression of stereotypes and prejudice, little is known about the brain structures involved in their acquisition. Here, we combined fMRI, a task involving learning the valence of different social groups, and modeling of the learning process involved in the development of biases in thinking about social groups that support prejudice. Participants read descriptions of valenced behaviors performed by members of novel social groups, with majority groups being more frequently encountered during learning than minority groups. A model-based fMRI analysis revealed that the anterior temporal lobe tracked the trial-by-trial changes in the valence associated with each group encountered in the task. Descriptions of behavior by group members that deviated from the group average (i.e., prediction errors) were associated with activity in the left lateral PFC, dorsomedial PFC, and lateral anterior temporal cortex. Minority social groups were associated with slower acquisition rates and more activity in the ventral striatum and ACC/dorsomedial PFC compared with majority groups. These findings provide new insights into the brain regions that (a) support the acquisition of prejudice and (b) detect situations in which an individual's behavior deviates from the prejudicial attitude held toward their group.

Oculomotor capture by stimuli that signal the availability of reward.

It is well known that eye movement patterns are influenced by both goal- and salience-driven factors. Recent studies, however, have demonstrated that objects that are nonsalient and task irrelevant can still capture our eyes if moving our eyes to those objects has previously produced reward. Here we demonstrate that training such an association between eye movements to an object and delivery of reward is not needed. Instead, an object that merely signals the availability of reward captures the eyes even when it is physically nonsalient and never relevant for the task. Furthermore, we show that oculomotor capture by reward is more reliably observed in saccades with short latencies. We conclude that a stimulus signaling high reward has the ability to capture the eyes independently of bottom-up physical salience or top-down task relevance and that the effect of reward affects early selection processes.

Location-based errors in change detection: A challenge for the slots model of visual working memory.

The slots model of visual working memory, despite its simplicity, has provided an excellent account of data across a number of change detection experiments. In the current research, we provide a new test of the slots model by investigating its ability to account for the increased prevalence of errors when there is a potential for confusion about the location in which items are presented during study. We assume that such location errors in the slots model occur when the feature information for an item in one location is swapped with the feature information for an item in another location. We show that such a model predicts two factors that will influence the extent to which location errors occur: (1) whether the test item changes to an "external" item not presented at study, or to an "internal" item presented at another location during study, and (2) the number of items in the study array. We manipulate these factors in an experiment, and show that the slots model with location errors fails to provide a satisfactory account of the observed data.

Delusions and prediction error: re-examining the behavioural evidence for disrupted error signalling in delusion formation.

INTRODUCTION: There is now significant evidence that prediction error signalling is mediated by dopamine in the midbrain, and that dopamine dysfunction is implicated in people experiencing psychotic symptoms, including delusions. There has also been significant theorizing and experimentation concerning the remaining link in this triad, namely that deviant prediction error signalling produces or maintains psychotic symptoms. METHODS: The research supporting the link between prediction error signalling and delusional symptoms was reviewed. Numerous studies indirectly support this link, but only one set of studies claim to directly test this hypothesis by combining three crucial elements: a patient sample, a manipulation of prediction error and neuroimaging. This particular set of studies were examined in detail. RESULTS: Important methodological limitations in these studies were observed, and a reinterpretation of their data was offered. CONCLUSIONS: Methodological inconsistencies significantly weaken the claims made by these studies, but their data are consistent with current theorizing and they are instructive for future lines of inquiry in this field.