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.

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 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.

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.

The influence of partner cues on the extinction of causal judgments in people.

Studies in laboratory animals have shown that the extinction of a conditioned stimulus, A, is regulated by the associative history of a second stimulus, X, when the two are extinguished in simultaneous compound: An inhibitory X protects A from extinction (Rescorla Learning & Behavior, 31, 124-132, 2003), whereas an excitatory X facilitates, and under some circumstances deepens, the extinction of A (Rescorla Journal of Experimental Psychology: Animal Behavior Processes, 26, 251-260, 2000, Journal of Experimental Psychology: Animal Behavior Processes, 32, 135-144, 2006). In the present study, we used the allergist task to examine whether the extinction of causal judgments in people is similarly regulated by the causal status of co-present stimuli. Experiment 1 showed that a cue trained as a conditioned inhibitor protected a target cue from extinction: The target extinguished in compound with the inhibitor was rated as being more causal of the outcome than was a target extinguished in compound with a control cue lacking inhibitory properties. In contrast, the remaining experiments showed that the extinction of a target cue was regulated by the presence, but not the causal status, of a partner cue: Target cues extinguished in compound were protected from extinction, and no evidence showed that an already extinguished partner conferred more protection (Exp. 2), or that an excitatory partner conferred any less protection (Exps. 2 and 3), or that an excitatory partner deepened the extinction of its already extinguished target. These findings are inconsistent with elemental models that rely on a common error term to explain associative changes in extinction. They are largely, but not completely, consistent with the configural model proposed by Pearce (Psychological Review, 94, 61-73, 1987), which predicts an ordering of levels of protection that was not observed.

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.

Absence of differential protection from extinction in human causal learning.

Elemental models of associative learning typically employ a common prediction-error term. Following a conditioning trial, they predict that the change in the strength of an association between a cue and an outcome is dependent upon how well the outcome was predicted. When multiple cues are present, they each contribute to that prediction. The same rule applies both to increases in associative strength during excitatory conditioning and the loss of associative strength during extinction. In five experiments using an allergy prediction task, we tested the involvement of a common error term in the extinction of causal learning. Two target cues were each paired with an outcome prior to undergoing extinction in compound either with a second excitatory cue or with a cue that had previously undergone extinction in isolation. At test, there was no difference in the causal ratings of the two target cues. Manipulations designed to bias participants toward elemental processing of cue compounds, to promote the acquisition of inhibitory associations, or to reduce generalization decrement between training and test were each without effect. These results are not consistent with common error term models of associative learning. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

An investigation of inattentional blindness using gaze and frequency tagging.

Inattentional blindness (IB) occurs when a salient object presented in plain sight goes unnoticed when its appearance is unexpected. Across two experiments, participants completed a classic dynamic IB task while eye movements and steady-state visual evoked potential (SSVEP) responses were continually recorded. This allowed us to measure the modulation of gaze and brain-based indices of attention during IB. While an SSVEP response to all stimuli including the unexpected object was attained, only gaze measures were able to discriminate noticers from nonnoticers. Experiment 1 used a prototypical sustained IB task and found that gaze toward the unexpected object was largely unrelated to noticing that object. Experiment 2 manipulated the contrast of the target and distractor stimuli, and instead observed a tight concordance between gazing at the unexpected object and reporting its presence. This task-based variability in gaze deployment is consistent with the broader literature and cumulatively delineates the challenges faced in translating lab-based IB research from the bench to the bedside. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

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.

A common error term regulates acquisition but not extinction of causal judgments in people.

In three experiments, we used the allergist task to examine the role of error correction mechanisms in the acquisition and extinction of causal judgments in people. Consistent with existing human and animal studies, acquisition of causal judgments was influenced by the discrepancy between the allergenic outcome and that predicted by all of the cues present on a trial (the "common error" term). However, in the present experiments, we failed to detect any evidence for the use of a common error term in extinction learning: Judgments of the allergenic properties of a cue were unaffected by the predictive value of the other cues present on a trial. This asymmetry in the use of a common error term in acquisition and extinction learning is inconsistent with previous animal studies and also with most models of associative learning. However, approaches that allow learning to be specific to a particular arrangement of elemental cues (context specific and state based) offer some explanation of the observed asymmetry.

Negative transfer in human associative learning.

Models of attentional allocation in associative learning are typically structured according to one of two guiding principles: the predictiveness principle, which posits that attention is paid to cues that have reliably predicted an outcome in the past, or the uncertainty principle, which states that attention is paid to cues about which little is known. Both principles are well supported by studies of animals. However, in studies of human learning, there is very little direct empirical support for the uncertainty principle. In the study reported here, we addressed this gap by investigating negative transfer, a phenomenon that may provide unique support for the uncertainty principle. In two human learning experiments using an allergist task, we replicated the primary findings of previous research on animal learning. We believe that these data provide the first direct evidence for the uncertainty principle in human associative learning.

Schizotypy is associated with difficulty maintaining multiple hypotheses.

Among neurocognitive accounts of delusions, there is a growing consensus that it is the certainty with which delusions are held, rather than their content that defines some beliefs as delusional. On a continuum model of psychosis, this inappropriate certainty ought to be present (albeit in an attenuated form) in healthy adults who score highly in schizotypy. It was hypothesised that this might be most evident in circumstances where the environment provides incomplete or probabilistic information, which thereby forces the participant to hold two imperfectly supported, concurrent hypotheses in mind. A cued visual search task was used to measure people's capacity to use partially predictive information (i.e., a cue that predicted the target may occur in one of the two locations) to facilitate speeded responding. As hypothesised, people's performance on the trials that required holding two hypotheses in mind concurrently was significantly and specifically associated with the positive components of schizotypy. This finding is consistent with a hyperfocusing of attention in schizophrenia, and may help explain why delusion-prone individuals have a tendency to "jump to conclusions" or be resistant to disconfirming information when faced with multiple, partially supported hypotheses.

Outcome unpredictability affects outcome-specific motivation to learn.

Outcome predictability effects in associative learning paradigms describe better learning about outcomes with a history of greater predictability in a similar but unrelated task compared with outcomes with a history of unpredictability. Inspired by the similarities between this phenomenon and the effect of uncontrollability in learned helplessness paradigms, here, we investigate whether learning about unpredictability decreases outcome-specific motivation to learn. We used a modified version of the allergy task, in which participants first observe the foods eaten by a fictitious patient, followed by allergic reactions that he subsequently suffers, some of which are perfectly predictable and others unpredictable. We then implemented an active learning method in a second task in which participants could only learn about either the previously predictable or unpredictable outcomes on each trial. At the beginning of each trial, participants had to decide whether they wanted to learn about one outcome category or the other. Participants at the beginning of the second task chose to learn about the previously predictable outcomes first and to learn about the previously unpredictable outcomes in later trials. This showed that unpredictability affects future motivation to learn in other circumstances. Interestingly, we did not find any sign of outcome predictability effect at the end of the second phase, suggesting that participants compensate for biased outcome sampling when making overt choices in ways that they may not when learning about both outcome categories simultaneously.

Reinforcement history shapes primary visual cortical responses: An SSVEP study.

Efficient learning requires allocating limited attentional resources to meaningful stimuli and away from irrelevant stimuli. This prioritization may occur via covert attention, evident in the activity of the visual cortex. We used steady-state visual evoked potentials (SSVEPs) to assess whether associability-driven changes in stimulus processing were evident in visuocortical responses. Participants were trained on a learned-predictiveness protocol, whereby one stimulus on each trial accurately predicted the correct response for that trial, and the other was irrelevant. In a second phase the task was arranged so that all cues were objectively predictive. Participants' overt attention (eye gaze) was affected by each cue's reinforcement history, as was their covert attention (SSVEP responses). These biases persisted into Phase 2 when all stimuli were objectively predictive, thereby demonstrating that learned attentional processes are evident in basic sensory processing, and exert an effect on covert attention above and beyond the effects of overt gaze bias.

The outcome predictability bias is evident in overt attention.

Previous studies of human associative learning have demonstrated that people's experience with a cueing stimulus will change how that cue is treated during subsequent learning. Typically, studies have shown that people pay more attention to cues that were informative in the past, and learn new information about these cues more rapidly (these cues are said to have a higher associability). It has recently been shown that to-be-predicted events (outcomes) can also differ in their associability as a consequence of prior experience. However, to date there is no direct evidence that this change in associability is accompanied by a change in attention, which would provide stronger evidence of a parallel with the effects observed previously with cueing stimuli. In two experiments, we examined this question by tracking eye-gaze to provide a measure of participants' overt attention, as they completed a cued visual search task in which outcome predictability was manipulated. The prior predictability of an outcome stimulus biased eye-gaze and learning rate, in a manner reminiscent of the gaze biases observed in tasks that manipulate cue associability. The present results support the view that outcomes, like cues, can vary in the degree to which they attract both attention and learning resources, as a function of their associative history. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Learned biases in the processing of outcomes: A brief review of the outcome predictability effect.

Much empirical work and theoretical discussion in the associative learning literature has focused on when and how a cue changes in its associability. A series of new findings in human learning preparations (collectively referred to as the "outcome predictability" effect) appear to show that outcomes vary in their capacity to enter into novel associations as a product of their associative history. This effect is reminiscent of how cues change in associability as a consequence of their reinforcement history. We review the new findings within a broader associative literature that has previously investigated how conditioning can modify the effectiveness of outcome events to motivate new learning. A variety of explanations arising from this review are then critically considered. The article concludes by identifying novel questions brought into focus by the outcome predictability effect. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Superstition predicts perception of illusory control.

Superstitions are common, yet we have little understanding of the cognitive mechanisms that bring them about. This study used a laboratory-based analogue for superstitious beliefs that involved people monitoring the relationship between undertaking an action (pressing a button) and an outcome occurring (a light illuminating). The task was arranged such that there was no objective contingency between pressing the button and the light illuminating - the light was just as likely to illuminate whether the button was pressed or not. Nevertheless, most people rated the causal relationship between the button press and the light illuminating to be moderately positive, demonstrating an illusion of causality. This study found that the magnitude of this illusion was predicted by people's level of endorsement of common superstitious beliefs (measured using a novel Superstitious Beliefs Questionnaire), but was not associated with mood variables or their self-rated locus of control. This observation is consistent with a more general individual difference or bias to overweight conjunctive events over disjunctive events during causal reasoning in those with a propensity for superstitious beliefs.

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.

Selective attention in human associative learning and recognition memory.

Four experiments examined the role of selective attention in a new causal judgment task that allowed measurement of both causal strength and cue recognition. In Experiments 1 and 2, blocking was observed; pretraining with 1 cue (A) resulted in reduced learning about a 2nd cue (B) when those 2 cues were trained in compound (AB+). Participants also demonstrated decreased recognition performance for the causally redundant Cue B, suggesting that less attention had been paid to it in training. This is consistent with the idea that attention is preferentially allocated toward the more predictive Cue A, and away from the less predictive Cue B (e.g., N. J. Mackintosh, 1975). Contrary to this hypothesis, in Experiments 3 and 4, participants demonstrated poorer recognition for the most predictive cues, relative to control cues. A new model, which is based on N. J. Mackintosh's (1975) model, is proposed to account for the observed relationship between the extent to which each cue is attended to, learned about, and later recognized