Violation of identity-specific action-effect prediction increases pupil size and attenuates auditory event-related potentials at P2 latencies when action-effects are behaviorally relevant.

Self-initiated sensory action effects are widely assumed to lead to less intense perception and reduced neural responses compared to externally triggered stimuli (sensory attenuation). However, it is unclear if sensory attenuation occurs in all cases of action-effect prediction. Specifically, when predicted action-effects are relevant to determine follow-up actions attenuation could be detrimental. We quantified auditory event-related potentials (ERP) in electroencephalography (EEG) when human participants created two-sound sequences by pressing two keys on a keyboard associated with different pitch, giving rise to identity-specific action-effect prediction after the first keypress. The first sound corresponded to (congruent) or violated (incongruent) the predicted pitch and was either relevant for the selection of the second keypress to correctly complete the sequence (Relevance) or irrelevant (Control Movement), or there was only one keypress and sound (Baseline). We found a diminished P2-timed ERP component in incongruent compared to congruent trials when the sound was relevant for the subsequent action. This effect of action-effect prediction was due to an ERP reduction for incongruent relevant sounds compared to incongruent irrelevant sounds at P2 latencies and correlated negatively with modulations of pupil dilation. Contrary to our expectation, we did not observe an N1 modulation by congruency in any condition. Attenuation of the N1 component seems absent for predicted identity-specific auditory action effects, while P2-timed ERPs as well as pupil size are sensitive to predictability, at least when action effects are relevant for the selection of the next action. Incongruent relevant stimuli thereby take a special place and seem to be subject to attentional modulations and error processing.

The topographical attenuation of cutaneous input is modulated at the ankle joint during gait.

The attenuation of sensory inputs via various methods has been demonstrated to impair balance control and alter locomotor behavior during human walking; however, the effects of attenuating foot sole sensation under distinct areas of the foot sole on lower extremity motor output remains poorly understood. Thus, the purpose of this study was to attenuate cutaneous feedback via regional hypothermia under five different areas of the foot sole and investigate the resultant modulation of kinematic and muscle activity during level walking. Electromyography from eight lower leg muscles, kinematics, and location of center of pressure was recorded from 48 healthy young adults completing walking trials with normal and reduced cutaneous sensation from bilateral foot soles. The results of this study highlight the modulatory response of the tibialis anterior in terminal stance (propulsion and toe-off) and medial gastrocnemius muscle throughout the entire stance phase of gait. The topographical organization of foot sole skin in response to the attenuation of cutaneous feedback from different areas of the foot sole significantly modified locomotor activity. Furthermore, the locomotor response to cutaneous attenuation under the same regions that we previously facilitated with tactile feedback do not oppose each other, suggesting different physiological changes to foot sole skin generate unique gait behaviors.

Exploring the internal forward model: action-effect prediction and attention in sensorimotor processing.

Action-effect predictions are believed to facilitate movement based on its association with sensory objectives and suppress the neurophysiological response to self- versus externally generated stimuli (i.e. sensory attenuation). However, research is needed to explore theorized differences in the use of action-effect prediction based on whether movement is uncued (i.e. volitional) or in response to external cues (i.e. stimulus-driven). While much of the sensory attenuation literature has examined effects involving the auditory N1, evidence is also conflicted regarding this component's sensitivity to action-effect prediction. In this study (n = 64), we explored the influence of action-effect contingency on event-related potentials associated with visually cued and uncued movement, as well as resultant stimuli. Our findings replicate recent evidence demonstrating reduced N1 amplitude for tones produced by stimulus-driven movement. Despite influencing motor preparation, action-effect contingency was not found to affect N1 amplitudes. Instead, we explore electrophysiological markers suggesting that attentional mechanisms may suppress the neurophysiological response to sound produced by stimulus-driven movement. Our findings demonstrate lateralized parieto-occipital activity that coincides with the auditory N1, corresponds to a reduction in its amplitude, and is topographically consistent with documented effects of attentional suppression. These results provide new insights into sensorimotor coordination and potential mechanisms underlying sensory attenuation.

More than fulfilled expectations: An electrophysiological investigation of varying cause-effect relationships and schizotypal personality traits as related to the sense of agency.

The sense of agency (SoA) is central to human experience. The comparator model, contrasting sensory prediction and action feedback, is influential but limited in explaining SoA. We investigated mechanisms beyond the comparator model, focusing on the processing of unpredictable stimuli, perimotor components of SoA, and their relation to schizotypy. ERPs were recorded from 18 healthy participants engaged in button-pressing tasks while perceiving tones with varying causal relationships with their actions. We investigated the processing of non-causally related tones, contrasted this to causally related tones, and examined perimotor correlates of subjective expectancy and experience of agency. We confirmed N100 attenuation for self-generated stimuli but found similar effects for expectancy-dependent processing of random tones. SoA also correlated with perimotor ERP components, modulated by schizotypy. Thus, neural processes preceding actions contribute to the formation of SoA and are associated with schizotypy. Unpredictable events also undergo sensory attenuation, implying additional mechanisms contributing to SoA.

Role of selective attention in fatigue in neurological disorders.

BACKGROUND: Chronic fatigue is a significant symptom in several diseases including traumatic and degenerative neurological disorders. While several studies have investigated the correlates of chronic fatigue, there is as yet no unifying framework to explain chronic fatigue. METHODS: In this narrative review, I investigate the role of selective attention in the development of chronic fatigue and discuss results within the framework of the sensory attenuation model of fatigue, which posits that fatigue is the phenomenological output of altered attention to sensory input. Following a short introduction of this framework, I present results from investigations that address attentional mechanisms in fatigue in multiple sclerosis, stroke, traumatic brain injury and Parkinson's disease. RESULTS: Attention was quantified in all four disease models using a variety of outcome measures, including behavioural, neurophysiological, structural and functional brain connectivity. The range of measures precluded direct comparison of results across disease conditions; however, in all four disease models there was evidence of poor selective attention that explained levels of chronic fatigue, supporting the sensory attenuation model of fatigue as a disease-independent mechanism of fatigue. Evidence was lacking to draw any conclusions about the direction of causality. CONCLUSION: The role of selective attention in development of fatigue is indicated. Future studies must focus on establishing causality and exploring attentional circuitry as a potential therapeutic target.

Temporal adaptation of sensory attenuation for self-touch.

The sensory consequences of our actions appear attenuated to us. This effect has been reported for external sensations that are evoked by auditory or visual events and for body-related sensations which are produced by self-touch. In the present study, we investigated the effects of prolonged exposure to a delay between an action and the generated sensation on sensory attenuation for self-touch. Previously, it has been shown that after being presented to a systematic exposure delay, artificially delayed touch can feel more intense and non-delayed touches can appear less intense. Here, we investigated the temporal spread of the temporal recalibration effect. Specifically, we wondered whether this temporal recalibration effect would affect only the delay that was used during exposure trials or if it would also modulate longer test delays. In the first two experiments, we tested three test delays (0, 100 and 400 ms) either in randomized or in blocked order. We found sensory attenuation in all three test intervals but no effect of the exposure delay. In Experiment 3, we replicated the experiment by Kilteni et al. (ELife 8:e42888, 2019. https://doi.org/10.7554/eLife.42888 ) and found evidence for temporal recalibration by exposure delay. Our data show that the temporal selectivity of sensory attenuation of self-touch depends on presenting a singular test delay only. Presenting multiple test delays leads to a temporally broad spread of sensory attenuation.

Virtual occlusion effects on the perception of self-initiated visual stimuli.

Virtual reality (VR) has established itself as a useful tool in the study of human perception in the laboratory. A recent study introduced a new approach to examine visual sensory attenuation (SA) effects in VR. Hand movements triggered the appearance of Gabor stimuli, which were either presented behind the participant's hand - not rendered in VR ("virtual occlusion") - or elsewhere on the display. Virtual occlusion led to a rightward shift of the psychometric curve, suggesting that self-generated hand movements reduced the perceived contrast of the stimulus. Since such attenuation effects might provide a window into the predictive processing of the sensory and cognitive apparatus, we sought to better understand the nature of the virtual occlusion effects. In our study, the presentation of test stimuli was either self-initiated, self-initiated with a variable delay, or triggered externally; the test stimuli were occluded or not. In conflict with our hypothesis, we found moderate to strong evidence for an absence of any horizontal shifts between the psychometric curves. However, virtual occlusion was associated with a decrease in the slope of the psychometric function. Our results suggest that virtual occlusion attenuated the relative perceptual sensitivity, so that participants had more difficulty discriminating contrast differences when the test stimulus was presented behind the hand. We tentatively conclude that, in the visual domain, the discriminability of stimulus intensity is modified by internal predictive cues (i.e., proprioception), possibly linked to shifts in covert spatial attention.

Methodological considerations for the force-matching task.

The force-matching task integrates haptic technology and electrical engineering to determine an individual's level of sensory attenuation to somatic stimuli. The task requires a detailed methodology to facilitate reliable and replicable estimates, and there has been a distinct lack of re-evaluation of the methodological processes related to this paradigm. In this task, participants are asked to match a force delivered to their finger, either by pressing directly on their own finger with their other hand (known as the direct condition) or by controlling the device using an external potentiometer to control the force indirectly through a torque motor (known as the slider condition). We analysed 138 participants to determine 1) the optimal number of replications (2, 4, 6, or 8 replications) of the target force, 2) the optimal time window (1-1.5 s, 1.5-2 s, 2-2.5 s and 2.5-3 s) to extract the estimate of sensory attenuation, 3) if participants' performance during the task improved, worsened or was stable across the experimental period regardless of condition, and 4) if learning effects were related to psychological traits. Results showed that the number of replications of the target forces may be reduced from 8 without compromising the estimate of sensory attenuation, the optimal time window for the extraction of the matched force is 2.5-3 s, the performance is stable over the duration of the experiment and not impacted by the measured psychological traits. In conclusion, we present a number of methodological considerations which improve the efficiency and reliability of the force-matching task. HIGHLIGHTS: • The force-matching task determines an individual's level of sensory attenuation • The optimal number of replications of the target force may be reduced from 8 • The optimal time window to extract the matched force is 2.5-3.0 s • The estimate of sensory attenuation is stable across the duration of the task.

Action-based predictions affect visual perception, neural processing, and pupil size, regardless of temporal predictability.

Sensory consequences of one's own action are often perceived as less intense, and lead to reduced neural responses, compared to externally generated stimuli. Presumably, such sensory attenuation is due to predictive mechanisms based on the motor command (efference copy). However, sensory attenuation has also been observed outside the context of voluntary action, namely when stimuli are temporally predictable. Here, we aimed at disentangling the effects of motor and temporal predictability-based mechanisms on the attenuation of sensory action consequences. During fMRI data acquisition, participants (N = 25) judged which of two visual stimuli was brighter. In predictable blocks, the stimuli appeared temporally aligned with their button press (active) or aligned with an automatically generated cue (passive). In unpredictable blocks, stimuli were presented with a variable delay after button press/cue, respectively. Eye tracking was performed to investigate pupil-size changes and to ensure proper fixation. Self-generated stimuli were perceived as darker and led to less neural activation in visual areas than their passive counterparts, indicating sensory attenuation for self-generated stimuli independent of temporal predictability. Pupil size was larger during self-generated stimuli, which correlated negatively with the blood oxygenation level dependent (BOLD) response: the larger the pupil, the smaller the BOLD amplitude in visual areas. Our results suggest that sensory attenuation in visual cortex is driven by action-based predictive mechanisms rather than by temporal predictability. This effect may be related to changes in pupil diameter. Altogether, these results emphasize the role of the efference copy in the processing of sensory action consequences.

Increased upper-limb sensory attenuation with age.

The pressure of our own finger on the arm feels differently than the same pressure exerted by an external agent: the latter involves just touch, whereas the former involves a combination of touch and predictive output from the internal model of the body. This internal model predicts the movement of our own finger, and hence the intensity of the sensation of the finger press is decreased. A decrease in intensity of the self-produced stimulus is called sensory attenuation. It has been reported that, because of decreased proprioception with age and an increased reliance on the prediction of the internal model, sensory attenuation is increased in older adults. In this study, we used a force matching paradigm to test whether sensory attenuation is also present over the arm and whether aging increases sensory attenuation. We demonstrated that, although both young and older adults overestimate a self-produced force, older adults overestimate it even more, showing an increased sensory attenuation. In addition, we also found that both younger and older adults self-produce higher forces when activating the homologous muscles of the upper limb. Although this is traditionally viewed as evidence for an increased reliance on internal model function in older adults because of decreased proprioception, proprioception appeared unimpaired in our older participants. This begs the question of whether an age-related decrease in proprioception is really responsible for the increased sensory attenuation observed in older people.NEW & NOTEWORTHY Forces generated externally (by the environment on the participant) and internally (by the participant on her/his body) are not perceived with the same intensity. Internally generated forces are perceived less intensely than externally generated ones. This difference in force sensation has been shown to be higher in elderly participants when the forces were applied on the fingers because of their impaired proprioception. Here we replicated this finding for the arm but suggest that it is unlikely to be linked to impaired proprioception.

Sensory attenuation from action observation.

A central claim of many embodied approaches to cognition is that understanding others' actions is achieved by covertly simulating the observed actions and their consequences in one's own motor system. If such a simulation occurs, it may be accomplished through forward models, a component of the motor system already known to perform simulations of actions and their consequences in order to support sensory-monitoring of one's own actions. Forward-model simulations cause an attenuation of sensory intensity, so if the simulations hypothesized by embodied cognition are indeed provided by forward models, then action observation should trigger this sensory attenuation. To test this hypothesis, the experiments reported here measured the perceived intensity of a touch sensation on the finger when participants observed an active touch (a finger reaching to touch a ball) vs. a passive touch (a ball rolling to touch an unmoving finger). The touch sensation was perceived as less intense during observation of active touch in comparison with observation of passive touch, providing evidence that forward models are indeed engaged during action observation. The strength of this sensory attenuation is compared and contrasted with a well-established sensory-amplification effect caused by visual attention. This sensory-amplification effect has not generally been considered in studies related to sensory attenuation in action observation, which may explain conflicting results reported in the field.

Dissociable effects of attention and expectation on perceptual sensitivity to action-outcomes.

Self-generated sensations evoke attenuated neural response - physiological attenuation - and is perceived with less intensity - perceptual attenuation. This phenomenon is referred as sensory attenuation and is proposed to reflect the silencing of predicted sensations. The present study aimed to investigate the independent contribution of expectation and attention on sensory attenuation. The expectation associated with the stimulus feature and the focus of attention was manipulated independently by orthogonal cues. We found pronounced sensory attenuation at the unattended location when the stimulus was self-generated (Experiment 1). When the stimulus was externally-generated (Experiment 2), sensory attenuation was observed at the attended location. Sensory attenuation of expected action-outcome was not observed when the attention cue was uninformative (Experiment 3A). The findings corroborate the claim from Bayesian models that attention mediates sensory attenuation. The results also highlight the paradoxes in Bayesian proposals of perception-action interaction.

Tactile motor attention induces sensory attenuation for sounds.

Sensory events appear reduced in intensity when we actively produce them. Here, we investigated sensory attenuation in a virtual reality setup that allowed us to manipulate the time of tactile feedback when pressing a virtual button. We asked whether tactile motor attention might shift to the tactile location that makes contact with the button. In experiment one, we found that a tactile impulse was perceived as more intense when button pressing. In a second experiment, participants pushed a button and estimated the intensity of sounds. We found sensory attenuation for sounds only when tactile feedback was provided at the time the movement goal was reached. These data indicate that attentional prioritization for the tactile modality during a goal-directed hand movement might lead to a transient reduction in sensitivity in other modalities, resulting in sensory attenuation for sounds.

I overthink-Therefore I am not: An active inference account of altered sense of self and agency in depersonalisation disorder.

This paper considers the phenomenology of depersonalisation disorder, in relation to predictive processing and its associated pathophysiology. To do this, we first establish a few mechanistic tenets of predictive processing that are necessary to talk about phenomenal transparency, mental action, and self as subject. We briefly review the important role of 'predicting precision' and how this affords mental action and the loss of phenomenal transparency. We then turn to sensory attenuation and the phenomenal consequences of (pathophysiological) failures to attenuate or modulate sensory precision. We then consider this failure in the context of depersonalisation disorder. The key idea here is that depersonalisation disorder reflects the remarkable capacity to explain perceptual engagement with the world via the hypothesis that "I am an embodied perceiver, but I am not in control of my perception". We suggest that individuals with depersonalisation may believe that 'another agent' is controlling their thoughts, perceptions or actions, while maintaining full insight that the 'other agent' is 'me' (the self). Finally, we rehearse the predictions of this formal analysis, with a special focus on the psychophysical and physiological abnormalities that may underwrite the phenomenology of depersonalisation.

Distinction of self-produced touch and social touch at cortical and spinal cord levels.

Differentiation between self-produced tactile stimuli and touch by others is necessary for social interactions and for a coherent concept of "self." The mechanisms underlying this distinction are unknown. Here, we investigated the distinction between self- and other-produced light touch in healthy volunteers using three different approaches: fMRI, behavioral testing, and somatosensory-evoked potentials (SEPs) at spinal and cortical levels. Using fMRI, we found self-other differentiation in somatosensory and sociocognitive areas. Other-touch was related to activation in several areas, including somatosensory cortex, insula, superior temporal gyrus, supramarginal gyrus, striatum, amygdala, cerebellum, and prefrontal cortex. During self-touch, we instead found deactivation in insula, anterior cingulate cortex, superior temporal gyrus, amygdala, parahippocampal gyrus, and prefrontal areas. Deactivation extended into brain areas encoding low-level sensory representations, including thalamus and brainstem. These findings were replicated in a second cohort. During self-touch, the sensorimotor cortex was functionally connected to the insula, and the threshold for detection of an additional tactile stimulus was elevated. Differential encoding of self- vs. other-touch during fMRI correlated with the individual self-concept strength. In SEP, cortical amplitudes were reduced during self-touch, while latencies at cortical and spinal levels were faster for other-touch. We thus demonstrated a robust self-other distinction in brain areas related to somatosensory, social cognitive, and interoceptive processing. Signs of this distinction were evident at the spinal cord. Our results provide a framework for future studies in autism, schizophrenia, and emotionally unstable personality disorder, conditions where symptoms include social touch avoidance and poor self-vs.-other discrimination.

The implicit sense of agency is not a perceptual effect but is a judgment effect.

The sense of agency (SoA) is characterized as the sense of being the causal agent of one's own actions, and it is measured in two forms: explicit and implicit. In the explicit SoA experiments, the participants explicitly report whether they have a sense of control over their actions or whether they or somebody else is the causal agent of seen actions; the implicit SoA experiments study how do participants' agentive or voluntary actions modify perceptual processes (like time, vision, tactility, and audition) without directly asking the participants to explicitly think about their causal agency or sense of control. However, recent implicit SoA literature reported contradictory findings of the relationship between implicit SoA reports and agency states. Thus, I argue that the purported implicit SoA reports are not agency-driven perceptual effects per se but are judgment effects, by showing that (a) the typical operationalizations in implicit SoA domain lead to perceptual uncertainty on the part of the participants, (b) under uncertainty, participants' implicit SoA reports are due to heuristic judgments which are independent of agency states, and (c) under perceptual certainty, the typical implicit SoA reports might not have occurred at all. Thus, I conclude that the instances of implicit SoA are judgments (or response biases)-under uncertainty-rather than perceptual effects.

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.

I'm a believer: Illusory self-generated touch elicits sensory attenuation and somatosensory evoked potentials similar to the real self-touch.

Sensory attenuation (i.e., the phenomenon whereby self-produced sensations are perceived as less intense compared to externally occurring ones) is among the neurocognitive processes that help distinguishing ourselves from others. It is thought to be rooted in the motor system (e.g., related to motor intention and prediction), while the role of body awareness, which necessarily accompanies any voluntary movement, in this phenomenon is largely unknown. To fill this gap, here we compared the perceived intensity, somatosensory evoked potentials, and alpha-band desynchronization for self-generated, other-generated, and embodied-fake-hand-generated somatosensory stimuli. We showed that sensory attenuation triggered by the own hand and by the embodied fake hand had the same behavioral and neurophysiological signatures (reduced subjective intensity, reduced of N140 and P200 SEP components and post-stimulus alpha-band desynchronization). Therefore, signals subserving body ownership influenced attenuation of somatosensory stimuli, possibly in a postdictive manner. This indicates that body ownership is crucial for distinguishing the source of the perceived sensations.

Expected TMS excites the motor system less effectively than unexpected stimulation.

The brain's response to sensory input is modulated by prediction. For example, sounds that are produced by one's own actions, or those that are strongly predicted by environmental cues, elicit an attenuated N1 component in the auditory evoked potential. It has been suggested that this form of sensory attenuation to stimulation produced by one's own actions is the reason we are unable to tickle ourselves. Here we examined whether the neural response to direct stimulation of the brain is attenuated by prediction in a similar manner. Transcranial magnetic stimulation (TMS) applied over primary motor cortex can be used to gauge the excitability of the motor system. Motor-evoked potentials (MEPs), elicited by TMS and measured in peripheral muscles, are larger when actions are being prepared and smaller when actions are voluntarily suppressed. We tested whether the amplitude of MEPs was attenuated under circumstances where the TMS pulse can be reliably predicted, even though control of the relevant motor effector was never required. Self-initiation of the TMS pulse and reliable cuing of the TMS pulse both produced attenuated MEP amplitudes, compared to those generated programmatically in an unpredictable manner. These results suggest that predictive coding may be governed by domain-general mechanisms responsible for all forms predictive learning. The findings also have important methodological implications for designing TMS experiments that control for the predictability of TMS pulses.

No evidence for somatosensory attenuation during action observation of self-touch.

The discovery of mirror neurons in the macaque brain in the 1990s triggered investigations on putative human mirror neurons and their potential functionality. The leading proposed function has been action understanding: Accordingly, we understand the actions of others by 'simulating' them in our own motor system through a direct matching of the visual information to our own motor programmes. Furthermore, it has been proposed that this simulation involves the prediction of the sensory consequences of the observed action, similar to the prediction of the sensory consequences of our executed actions. Here, we tested this proposal by quantifying somatosensory attenuation behaviourally during action observation. Somatosensory attenuation manifests during voluntary action and refers to the perception of self-generated touches as less intense than identical externally generated touches because the self-generated touches are predicted from the motor command. Therefore, we reasoned that if an observer simulates the observed action and, thus, he/she predicts its somatosensory consequences, then he/she should attenuate tactile stimuli simultaneously delivered to his/her corresponding body part. In three separate experiments, we found a systematic attenuation of touches during executed self-touch actions, but we found no evidence for attenuation when such actions were observed. Failure to observe somatosensory attenuation during observation of self-touch is not compatible with the hypothesis that the putative human mirror neuron system automatically predicts the sensory consequences of the observed action. In contrast, our findings emphasize a sharp distinction between the motor representations of self and others.