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Kilteni, K.

Publications and source records attributed to Kilteni, K..

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Motor prediction reduces beta-band power and enhances cerebellar-somatosensory connectivity before self-touch to enable its attenuation

Motor control theories suggest that the brain uses forward models to predict self-generated tactile input during voluntary movements, thereby reducing the intensity of reafferent tactile sensations. When ones own body is the target, this phenomenon is called self-touch attenuation. Although self-touch attenuation is well-documented, it remains unclear how prediction-related neural mechanisms drive attenuation before the self-touch input. We used magnetoencephalography (MEG) to examine the neural correlates of self-touch prediction. 24 human participants (12 female, 12 male) performed a self-touch, and two control tasks. In one control, they received externally generated touch without movement. In the other, the touch was triggered by the participants movement, but the hands were spatially misaligned. This manipulation is known to weaken attenuation despite identical tactile input, movement, and task demands, because the sensorimotor context reduces prediction of touch at that body site. Self-touch evoked weaker somatosensory activity (M50 component) than both control conditions. A psychophysics task mirrored the pattern of neural attenuation, as the perception of self-touch was attenuated compared to the two control conditions. To isolate predictive neural mechanisms from general movement-related activity, we subtracted activity from corresponding stimulus-absent trials. Comparing self-touch with misaligned touch allowed us to refine the signal specific to predictive processing in self-touch and revealed greater pre-stimulus beta-band desynchronization and increased cerebellar-to-somatosensory connectivity before self-touch compared to misaligned touch. Our results provide the first evidence of predictive neural activity that shapes the sensory consequences of self-touch, offering insights into the mechanisms through which predictive models modulate somatosensory processing. Significance statementThe brain is thought to predict and attenuate the sensory consequences of self-generated actions, but neural evidence for prediction before sensation has been limited. Using magnetoencephalography, we show that self-touch attenuation is preceded by beta-band desynchronization and increased directed connectivity from the cerebellum to the primary somatosensory cortex. These effects cannot be attributed to movement, as they were reduced in a control condition with similar motor output but lower congruence between the action and tactile consequence, suggesting spatially specific predictive processing. Our study provides the first neural evidence of cerebellar influence on cortical sensory areas before self-touch. These pre-stimulus effects support forward models of sensorimotor control and shed new light on how the brain anticipates and modulates upcoming sensory input.

neuroscience↗

Predictions of bimanual self-touch determine the temporal tuning of somatosensory perception

We effortlessly distinguish between touching ourselves with our hands and being touched by other people or objects. Motor control theories posit that this distinction is made possible by the brain predicting the somatosensory consequences of our voluntary movements based on an efference copy, and attenuating our responses to the predicted self-touch. However, it remains unclear how these predictions impact somatosensory perception at times other than during self- touch: for example, as our hand reaches to touch our body or moves away from it. Here participants discriminated forces applied on their left index finger by a motor. The forces were applied during the reaching movement of their right hand towards the left hand, including the time the reaching ended by simulating self-touch between the hands, or after the reaching movement. We observed that the forces on the left hand felt progressively weaker during the reaching phase, reached their minimum perceived intensity at the time of self-touch, and quickly recovered after the end of the reaching. All effects were replicated with a new cohort of participants that further demonstrated that this gradual attenuation of the perceived magnitude of touch vanished during similar right hand reaching movements that did not produce expectations for self-touch between the two hands. Together, our results indicate a temporal tuning of somatosensory perception during movements to self-touch and underscore the role of sensorimotor context in forming predictions that attenuate the intensity of self- generated touch.

neuroscience↗

Divisive attenuation based on noisy sensorimotor predictions accounts for excess variability in self-touch

When one part of the body exerts force on another, the resulting tactile sensation is perceived as weaker than when the same force is applied by an external agent. This phenomenon has been studied using a force matching task, in which observers are first exposed to an external force on a passive finger and then instructed to reproduce the sensation by directly pressing on the passive finger with a finger of the other hand: healthy participants consistently exceed the original force level. However, this exaggeration of the target force is not observed if the observer generates the matching force indirectly, by adjusting a joystick or slider that controls the force output of a motor. Here we present the first detailed computational account of the processes leading to exaggeration of target forces in the force matching task, incorporating attenuation of sensory signals based on motor predictions. The model elucidates previously unappreciated contributions of multiple sources of noise, including memory noise, in determining matching force output, and shows that quantifying attenuation as the discrepancy between direct and indirect self-generated forces isolates its predictive component. Our computational account makes the prediction that attenuated sensations will display greater trial-to-trial variability than unattenuated ones, because they incorporate additional noise from motor prediction. Quantitative model fitting of new and existing force matching data confirmed the prediction of excess variability in self-generated forces and provided evidence for a divisive rather than subtractive mechanism of attenuation, while highlighting its predictive nature. NEW & NOTEWORTHYWe formulate a detailed computational account of sensory attenuation in force matching tasks that disambiguates contributions of perceptual, memory and prediction noise in order to isolate a pure measure of attenuation strength. Analysis of data from nearly 500 participants shows that attenuated sensations display increased trial-to-trial variability, consistent with incorporating additional noise inherent to motor prediction. These results support a divisive - rather than subtractive - reduction in the sensation of self-generated forces based on predicted reafference.

neuroscience↗

Dynamic changes in somatosensory and cerebellar activity mediate temporal recalibration of self-touch

An organisms ability to accurately anticipate the sensations caused by its own actions is crucial for a wide range of behavioral, perceptual, and cognitive functions. Notably, the sensorimotor expectations produced when touching ones own body attenuate such sensations, making them feel weaker and less ticklish and rendering them easily distinguishable from potentially harmful touches of external origin. How the brain learns and keeps these action-related sensory expectations updated is unclear. We employed psychophysics and functional magnetic resonance imaging to pinpoint the behavioral and neural substrates of dynamic recalibration of expected temporal delays in self-touch. Psychophysical results revealed that self-touches were less attenuated after systematic exposure to delayed self-generated touches, while responses in the contralateral somatosensory cortex that normally distinguish between delayed and nondelayed self-generated touches became indistinguishable. During the exposure, the ipsilateral anterior cerebellum showed increased activity, supporting its proposed role in recalibrating sensorimotor predictions. Moreover, responses in the cingulate areas gradually increased, suggesting that as delay adaptation progresses, the nondelayed self-touches trigger activity related to cognitive conflict. Together, our results show that sensorimotor predictions in the simplest act of touching ones own body are upheld by a sophisticated and flexible neural mechanism that maintains them accurate in time.

neuroscience↗

Aging exerts a limited influence on the perception of self-generated and externally generated touch

Touch generated by our voluntary movements is attenuated both at the perceptual and neural level compared to touch of the same intensity delivered to our body by another person or machine. This somatosensory attenuation phenomenon relies on the integration of somatosensory input and predictions about the somatosensory consequences of our actions. Previous studies have reported increased somatosensory attenuation in elderly people, proposing an overreliance on sensorimotor predictions to compensate for age-related declines in somatosensory perception; however, recent results have challenged this direct relationship. In a preregistered study, we used a force-discrimination task to assess whether aging increases somatosensory attenuation and whether this increase is explained by decreased somatosensory precision in elderly individuals. Although 94% of our sample (n = 108, 21-77 years old) perceived their self-generated touches as weaker than externally generated touches of identical intensity (somatosensory attenuation) regardless of age, we did not find a significant increase in somatosensory attenuation in our elderly participants (65-77 years old), but a trend when considering only the oldest subset (69-77 years old). Moreover, we did not observe a significant age-related decline in somatosensory precision or a significant relationship of age with somatosensory attenuation. Together, our results suggest that aging exerts a limited influence on the perception of self-generated and externally generated touch and indicate a less direct relationship between somatosensory precision and attenuation in the elderly individuals than previously proposed. New and NoteworthySelf-generated touch is attenuated compared to externally generated touch of identical intensity. This somatosensory attenuation has been previously shown to be increased in elderly participants, but it remains unclear whether it is related to age-related somatosensory decline. In our preregistered study, we observed a trend for increased somatosensory attenuation in our oldest participants ([≥]69 years), but we found no evidence of an age-related decline in somatosensory function or a relationship of age with somatosensory attenuation.

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Action does not enhance but attenuates predicted touch

Dominant motor control theories propose that the brain predicts and attenuates the somatosensory consequences of actions, referred to as somatosensory attenuation. Support comes from psychophysical and neuroimaging studies showing that touch applied on a passive hand elicits attenuated perceptual and neural responses if it is actively generated by ones other hand, compared to identical touch from an external origin. However, recent experimental findings have challenged this view by providing psychophysical evidence that the perceived intensity of touch on the passive hand is enhanced if the active hand does not receive touch simultaneously with the passive hand (somatosensory enhancement) and by further attributing attenuation to the double tactile stimulation of the hands upon contact. Here, we directly contrasted the hypotheses of the attenuation and enhancement models regarding how action influences somatosensory perception by manipulating whether the active hand contacts the passive hand. We further assessed somatosensory perception in the absence of any predictive cues in a condition that turned out to be essential for interpreting the experimental findings. In three preregistered experiments, we demonstrate that action does not enhance the predicted touch (Experiment 1), that the previously reported "enhancement" effects are driven by the reference condition used (Experiment 2), and that self-generated touch is robustly attenuated regardless of whether the two hands make contact (Experiment 3). Our results provide conclusive evidence that action does not enhance but attenuates predicted touch and prompt a reappraisal of recent experimental findings upon which theoretical frameworks proposing a perceptual enhancement by action prediction are based. HighlightsO_LIDominant motor control theories propose that action attenuates or cancels predicted touch. C_LIO_LIRecent theories propose that action enhances predicted touch. C_LIO_LIWe show that action does not enhance but attenuates predicted touch. C_LI

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Brief temporal perturbations in somatosensory reafference disrupt perceptual and neural attenuation and increase supplementary motor-cerebellar connectivity

Intrinsic delays in sensory feedback can be detrimental for motor control. As a compensation strategy, the brain predicts the sensory consequences of movement via a forward model on the basis of a copy of the motor command. Using these predictions, the brain attenuates the somatosensory reafference to facilitate the processing of exafferent information. Theoretically, this predictive attenuation gets disrupted by (even minimal) temporal errors between the predicted and the actual reafference, but direct evidence for such disruption is lacking since previous neuroimaging studies contrasted conditions of nondelayed reafferent input with exafferent one. Here, we combined psychophysics with functional magnetic resonance imaging to test whether subtle perturbations in the timing of somatosensory reafference disrupt its predictive processing. Twenty-eight participants generated touches on their left index finger by tapping a sensor with their right index finger. The touches on the left index finger were delivered at the time of the two fingers contact or with a 100 ms delay. We found that such brief temporal perturbations disrupted the attenuation of the somatosensory reafference both at the perceptual and neural level, leading to greater somatosensory and cerebellar responses and weaker somatosensory connectivity with the cerebellum proportionally to perceptual changes. Moreover, we observed increased connectivity of the supplementary motor area with the cerebellum during the perturbations. We interpret these effects as the failure of the forward model to predictively attenuate the delayed somatosensory reafference and the return of the prediction error to the motor centers, respectively. Significance statementOur brain receives the somatosensory feedback of our movements with delay. To counteract these delays, motor control theories postulate that the brain predicts the timing of the somatosensory consequences of our movements and attenuates sensations received at that timing. This makes a self-generated touch feel weaker than an identical external touch. However, how subtle temporal errors between the predicted and the actual somatosensory feedback perturb this predictive attenuation remains unknown. We show that such errors make the otherwise attenuated touch feel stronger, elicit stronger somatosensory responses, weaken the cerebellar connectivity with somatosensory areas, and increase it with motor areas. These findings show that motor and cerebellar areas are fundamental in forming temporal predictions about the sensory consequences of our movements.

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The positive dimension of schizotypy is associated with a reduced attenuation and precision of self-generated touch

The brain predicts the sensory consequences of our movements and uses these predictions to attenuate the perception of self-generated sensations. Accordingly, self-generated touch feels weaker than externally generated touch of identical intensity. In schizophrenia, this somatosensory attenuation is substantially reduced, suggesting that patients with positive symptoms fail to accurately predict and process self-generated touch. Here we hypothesized that a similar impairment might exist in healthy nonclinical individuals with high positive schizotypal traits. One hundred healthy participants (53 female) scored for schizotypal traits and underwent a well-established psychophysics force discrimination task to quantify how they perceived self-generated and externally generated touch. The perceived intensity of tactile stimuli delivered to their left index finger (magnitude) and the ability to discriminate the stimuli (precision) were measured. We observed that higher positive schizotypal traits were associated with reduced somatosensory attenuation and poorer somatosensory precision of self-generated touch. These effects were specific to positive schizotypy and were not observed for the negative or disorganized dimensions of schizotypy. The results suggest that positive schizotypal traits are associated with a reduced ability to predict and process self-generated tactile stimuli. Given that the positive dimension of schizotypy represents the analogue of positive psychotic symptoms of schizophrenia, deficits in processing self-generated tactile information could indicate increased liability to schizophrenia.

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No evidence for somatosensory attenuationduring 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 programs. 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 behaviorally 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 simulates the observed action. In contrast, our findings emphasize a sharp distinction between the motor representations of self and others.

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Predictive attenuation of touch and tactile gating are distinct perceptual phenomena

In recent decades, research on somatosensory perception has led to two important observations. First, self-generated touches that are predicted by voluntary movements become attenuated compared to externally generated touches of the same intensity (attenuation). Second, externally generated touches feel weaker and are more difficult to detect during movement than at rest (gating). Currently, researchers often consider gating and attenuation the same suppression process; however, this assumption is unwarranted because, despite more than forty years of research, no study has combined them in a single paradigm. We quantified how people perceive self-generated and externally generated touches during movement and rest. We show that whereas voluntary movement gates the precision of both self-generated and externally generated touch, the amplitude of self-generated touch is robustly attenuated compared to externally generated touch. Furthermore, attenuation and gating do not interact and are not correlated, and we conclude that they represent distinct perceptual phenomena.

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Functional connectivity between the cerebellum and somatosensory areas implements the attenuation of self-generated touch

Since the early 1970s, numerous behavioral studies have shown that self-generated touch feels less intense than the same touch applied externally. Computational motor control theories have suggested that cerebellar internal models predict the somatosensory consequences of our movements and that these predictions attenuate the perception of the actual touch. Despite this influential theoretical framework, little is known about the neural basis of this predictive attenuation. This is due to the limited number of neuroimaging studies, the presence of conflicting results about the role and the location of cerebellar activity, and the lack of behavioral measures accompanying the neural findings. Here, we combined psychophysics with functional magnetic resonance imaging to detect the neural processes underlying somatosensory attenuation in male and female healthy human participants. Activity in bilateral secondary somatosensory areas was attenuated when the touch was presented during a self-generated movement (self-generated touch) than in the absence of movement (external touch). An additional attenuation effect was observed in the cerebellum that is ipsilateral to the passive limb receiving the touch. Importantly, we further found that the degree of functional connectivity between the ipsilateral cerebellum and the contralateral primary and bilateral secondary somatosensory areas was linearly and positively related to the degree of behaviorally assessed attenuation; that is, the more participants perceptually attenuated their self-generated touches, the stronger this corticocerebellar coupling. Collectively, these results suggest that the ipsilateral cerebellum is fundamental in predicting self-generated touch and that this structure implements somatosensory attenuation via its functional connectivity with somatosensory areas.\n\nSignificance statementWhen we touch our hand with the other, the resulting sensation feels less intense than when another person or a machine touches our hand with the same intensity. Early computational motor control theories have proposed that the cerebellum predicts and cancels the sensory consequences of our movements; however, the neural correlates of this cancelation remain unknown. By means of functional magnetic resonance imaging, we show that the more participants attenuate the perception of their self-generated touch, the stronger the functional connectivity between the cerebellum and the somatosensory cortical areas. This provides conclusive evidence about the role of the cerebellum in predicting the sensory feedback of our movements and in attenuating the associated percepts via its connections to early somatosensory areas.

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Efference copy is necessary for the attenuation of self-generated touch

A self-generated touch feels less intense than an external touch of the exact same intensity. According to a prevalent computational theory of motor control, this attenuation occurs because the brain uses internal forward models to predict the somatosensory consequences of our movements using a copy of the motor command, i.e., the efference copy. These tactile predictions are then used to suppress the perceived intensity of the actual tactile feedback. Despite being highly influential, the core assumption of theory has never been tested; that is, whether the efference copy is necessary for somatosensory attenuation. A possible alternative hypothesis is that a predictable contact of two of ones own body parts is sufficient. Using a psychophysical task, we quantified the attenuation of touch applied on the participants left index finger when the touch was triggered by the active or passive movement of the participants right index finger and when it was externally generated in the absence of any movement. We observed somatosensory attenuation only when the touch was triggered by the voluntary movement of the participants finger. In contrast, during the passive movement, the intensity of the touch was perceived to be as strong as when the touch was externally triggered. In both active and passive movement conditions, the participants showed the same discrimination capacity. Electromyographic recordings confirmed minimal activity of the right hand during the passive movement. Together, our results suggest that the efference copy is necessary for computing the somatosensory predictions that produce the attenuation of self-generated touch.

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Rapid learning and unlearning of predicted sensory delays in self-generated touch

Self-generated touch feels less intense and less ticklish than identical externally generated touch. This somatosensory attenuation occurs because the brain predicts the tactile consequences of our self-generated movements. To produce attenuation, the tactile predictions need to be time-locked to the movement, but how the brain maintains this temporal tuning remains unknown. Using a bimanual self-touch paradigm, we demonstrate that people can rapidly unlearn to attenuate touch immediately after their movement and learn to attenuate delayed touch instead, after repeated exposure to a systematic delay between the movement and the resulting touch. The magnitudes of the unlearning and learning effects are correlated and dependent on the number of trials that participants have been exposed to. We further show that delayed touches feel less ticklish and non-delayed touches more ticklish after exposure to the systematic delay. These findings demonstrate that the attenuation of self-generated touch is adaptive.

neuroscience↗

Postdoc X-ray in Europe 2017: work conditions, productivity, institutional support and career outlooks

1.Executive SummaryThis survey and data analysis were conducted by the European Network of Postdoctoral Associations (ENPA) with the aim of assessing the current research and work conditions, aspirations and support received by postdoctoral researchers working in Europe. The Results section is structured into three main parts. The first one describes the study sample of European postdoctoral researchers, including participants demographics, funding sources and income, research outputs, and teaching opportunities. The second section focuses on their professional aspirations and institutional support provided. The third part describes the level of engagement of postdoctoral researchers and their institutions in working towards better research conditions and career development, and what initiatives are emerging within this community. Our Conclusions section pulls together this comprehensive analysis, highlighting some of the most concerning issues currently affecting postdoctoral researchers in Europe. We also make a number of recommendations that would significantly improve the career expectations and aspirations of postdoctoral researchers. These are listed below. Conclusions and recommendationsO_LILonger postdoctoral periods in Southern Europe despite higher publication metrics Recommendation Institutions in Southern Europe should develop clear criteria to support postdoctoral researchers career progression. C_LIO_LISouthern and Eastern Europe pay the lowest salaries and have the lowest number of foreign postdoctoral researchers Recommendation The salary differences across European countries should be addressed as this could be a barrier to mobility and knowledge exchange from higher to lower pay regions. C_LIO_LILack of access to funding is a significant concern of postdoctoral researchers Recommendation Discrepancies in access to funding should be minimized across the different European areas. C_LIO_LIPostdoctoral researchers in Europe work longer hours than required by contract Recommendation The culture of overwork in the research environment should be addressed in order to protect researchers against the risks associated with long hours at work. C_LIO_LIThe majority of full-time postdoctoral work contracts includes an exclusivity clause Recommendation Inclusion of exclusivity clauses in contracts for postdoctoral researchers should be optional in order to allow them to enhance their employability outside academia. C_LIO_LIPostdoctoral researchers career development is poorly supported by their institutions Recommendation Postdoctoral researchers career prospects and career management should be much more supported by institutions in coordination with postdoctoral associations. C_LIO_LILack of postdoctoral representation in governance is linked to unclear institutional duties and rights Recommendation Institutional governance bodies should include postdoctoral researcher representatives. This would ensure that the views of this vital staff group are heard, as well as making postdoctoral researchers feel more engaged with their own institutions. A flexible and proactive communication strategy at the institution and research group level should be developed, taking into account the sometimes transient nature of postdoctoral researchers posts. C_LIO_LIResearchers show higher engagement with their local postdoctoral associations than with workers unions Recommendation Postdoctoral associations are an essential way to advocate for postdoctoral researchers at the governance level. Institutions should engage with, promote and support the work of postdoctoral associations. C_LI

scientific communication and education↗