The effect of self-generated versus externally generated actions on timing, duration, and amplitude of blood oxygen level dependent response for visual feedback processing.

It has been widely assumed that internal forward models use efference copies to create predictions about the sensory consequences of our own actions. While these predictions have frequently been associated with a reduced blood oxygen level dependent (BOLD) response in sensory cortices, the timing and duration of the hemodynamic response for the processing of video feedback of self-generated (active) versus externally generated (passive) movements is poorly understood. In the present study, we tested the hypothesis that predictive mechanisms for self-generated actions lead to early and shorter neural processing compared with externally generated movements. We investigated active and passive movements using a custom-made fMRI-compatible movement device. Visual video feedback of the active and passive movements was presented in real time or with variable delays. Participants had to judge whether the feedback was delayed. Timing and duration of BOLD impulse response was calculated using a first (temporal derivative [TD]) and second-order (dispersion derivative [DD]) Taylor approximation. Our reanalysis confirmed our previous finding of reduced BOLD response for active compared to passive movements. Moreover, we found positive effects of the TD and DD in the supplementary motor area, cerebellum, visual cortices, and subcortical structures, indicating earlier and shorter hemodynamic responses for active compared to passive movements. Furthermore, earlier activation in the putamen for active compared to passive conditions was associated with reduced delay detection performance. These findings indicate that efference copy-based predictive mechanisms enable earlier processing of action feedback, which might have reduced the ability to detect short delays between action and feedback.

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.

Non-stimulated regions in early visual cortex encode the contents of conscious visual perception.

Predictions shape our perception. The theory of predictive processing poses that our brains make sense of incoming sensory input by generating predictions, which are sent back from higher to lower levels of the processing hierarchy. These predictions are based on our internal model of the world and enable inferences about the hidden causes of the sensory input data. It has been proposed that conscious perception corresponds to the currently most probable internal model of the world. Accordingly, predictions influencing conscious perception should be fed back from higher to lower levels of the processing hierarchy. Here, we used functional magnetic resonance imaging and multivoxel pattern analysis to show that non-stimulated regions of early visual areas contain information about the conscious perception of an ambiguous visual stimulus. These results indicate that early sensory cortices in the human brain receive predictive feedback signals that reflect the current contents of conscious perception.

Different contributions of efferent and reafferent feedback to sensorimotor temporal recalibration.

Adaptation to delays between actions and sensory feedback is important for efficiently interacting with our environment. Adaptation may rely on predictions of action-feedback pairing (motor-sensory component), or predictions of tactile-proprioceptive sensation from the action and sensory feedback of the action (inter-sensory component). Reliability of temporal information might differ across sensory feedback modalities (e.g. auditory or visual), which in turn influences adaptation. Here, we investigated the role of motor-sensory and inter-sensory components on sensorimotor temporal recalibration for motor-auditory (button press-tone) and motor-visual (button press-Gabor patch) events. In the adaptation phase of the experiment, action-feedback pairs were presented with systematic temporal delays (0 ms or 150 ms). In the subsequent test phase, audio/visual feedback of the action were presented with variable delays. The participants were then asked whether they detected a delay. To disentangle motor-sensory from inter-sensory component, we varied movements (active button press or passive depression of button) at adaptation and test. Our results suggest that motor-auditory recalibration is mainly driven by the motor-sensory component, whereas motor-visual recalibration is mainly driven by the inter-sensory component. Recalibration transferred from vision to audition, but not from audition to vision. These results indicate that motor-sensory and inter-sensory components contribute to recalibration in a modality-dependent manner.

Commonalities and differences in predictive neural processing of discrete vs continuous action feedback.

Sensory action consequences are highly predictable and thus engage less neural resources compared to externally generated sensory events. While this has frequently been observed to lead to attenuated perceptual sensitivity and suppression of activity in sensory cortices, some studies conversely reported enhanced perceptual sensitivity for action consequences. These divergent findings might be explained by the type of action feedback, i.e., discrete outcomes vs. continuous feedback. Therefore, in the present study we investigated the impact of discrete and continuous action feedback on perceptual and neural processing during action feedback monitoring. During fMRI data acquisition, participants detected temporal delays (0-417 ms) between actively or passively generated wrist movements and visual feedback that was either continuously provided during the movement or that appeared as a discrete outcome. Both feedback types resulted in (1) a neural suppression effect (active

Neural Correlates of Self-other Distinction in Patients with Schizophrenia Spectrum Disorders: The Roles of Agency and Hand Identity.

Schizophrenia spectrum disorders (SSD) are characterized by disturbed self-other distinction. While previous studies associate abnormalities in the sense of agency (ie, the feeling that an action and the resulting sensory consequences are produced by oneself) with disturbed processing in the angular gyrus, passive movement conditions to isolate contributions of motor predictions are lacking. Furthermore, the role of body identity (ie, visual features determining whether a seen body part belongs to oneself) in self-other distinction is unclear. In the current study, fMRI was used to assess the roles of agency and hand identity in self-other distinction. Patients with SSD and healthy controls (HC) performed active and passive hand movements (agency manipulation) while seeing their own or someone else's hand moving in accordance with their action (hand identity manipulation). Variable delays (0-417 ms) between movement and feedback had to be detected. Our results showed overall lower delay detection performances during active than passive conditions; however, these differences were reduced in patients when the own hand was displayed. On a neural level, we found that in HC, activation in the right angular gyrus was modulated by agency and hand identity. In contrast, agency and hand identity revealed no overlapping activation in patients, due to reduced effects of agency. Importantly, HC and SSD patients shared similar effects of hand identity in the angular gyrus. Our results suggest that disturbances of self-other distinction in SSD are particularly driven by agency, while self-other distinction based on hand identity might be spared.

Seeing your own or someone else's hand moving in accordance with your action: The neural interaction of agency and hand identity.

Forward models can predict sensory consequences of self-action, which is reflected by less neural processing for actively than passively generated sensory inputs (BOLD suppression effect). However, it remains open whether forward models take the identity of a moving body part into account when predicting the sensory consequences of an action. In the current study, fMRI was used to investigate the neural correlates of active and passive hand movements during which participants saw either an on-line display of their own hand or someone else's hand moving in accordance with their movement. Participants had to detect delays (0-417 ms) between their movement and the displays. Analyses revealed reduced activation in sensory areas and higher delay detection thresholds for active versus passive movements. Furthermore, there was increased activation in the hippocampus, the amygdala, and the middle temporal gyrus when someone else's hand was seen. Most importantly, in posterior parietal (angular gyrus and precuneus), frontal (middle, superior, and medial frontal gyrus), and temporal (middle temporal gyrus) regions, suppression for actively versus passively generated feedback was stronger when participants were viewing their own compared to someone else's hand. Our results suggest that forward models can take hand identity into account when predicting sensory action consequences.

Transcranial direct current stimulation improves action-outcome monitoring in schizophrenia spectrum disorder.

Patients with schizophrenia spectrum disorder often demonstrate impairments in action-outcome monitoring. Passivity phenomena and hallucinations, in particular, have been related to impairments of efference copy-based predictions which are relevant for the monitoring of outcomes produced by voluntary action. Frontal transcranial direct current stimulation has been shown to improve action-outcome monitoring in healthy subjects. However, whether transcranial direct current stimulation can improve action monitoring in patients with schizophrenia spectrum disorder remains unknown. We investigated whether transcranial direct current stimulation can improve the detection of temporal action-outcome discrepancies in patients with schizophrenia spectrum disorder. On 4 separate days, we applied sham or left cathodal/right anodal transcranial direct current stimulation in a randomized order to frontal (F3/F4), parietal (CP3/CP4) and frontoparietal (F3/CP4) areas of 19 patients with schizophrenia spectrum disorder and 26 healthy control subjects. Action-outcome monitoring was assessed subsequent to 10 min of sham/transcranial direct current stimulation (1.5 mA). After a self-generated (active) or externally generated (passive) key press, subjects were presented with a visual outcome (a dot on the screen), which was presented after various delays (0-417 ms). Participants had to detect delays between the key press and the visual consequence. Symptom subgroups were explored based on the presence or absence of symptoms related to a paranoid-hallucinatory syndrome. In general, delay-detection performance was impaired in the schizophrenia spectrum disorder compared to the healthy control group. Interaction analyses showed group-specific (schizophrenia spectrum disorder versus healthy control group) and symptom-specific (with/without relevant paranoid-hallucinatory symptoms) transcranial direct current stimulation effects. Post hoc tests revealed that frontal transcranial direct current stimulation improved the detection of long delays in active conditions and reduced the proportion of false alarms in undelayed trials of the passive condition in patients. The patients with no or few paranoid-hallucinatory symptoms benefited especially from frontal transcranial direct current stimulation in active conditions, while improvement in the patients with paranoid-hallucinatory symptoms was predominantly reflected in reduced false alarm rates in passive conditions. These data provide some first evidence for the potential utility of transcranial direct current stimulation in improving efference copy mechanisms and action-outcome monitoring in schizophrenia spectrum disorder. Current data indicate that improving efference copy-related processes can be especially effective in patients with no or few positive symptoms, while intersensory matching (i.e. task-relevant in passive conditions) could be more susceptible to improvement in patients with paranoid-hallucinatory symptoms.

Predictive perception of self-generated movements: Commonalities and differences in the neural processing of tool and hand actions.

Tool use is one of the most remarkable skills of the human species, enabling complex interactions with the environment. To establish such interactions, we predict the sensory consequences of our actions based on a copy of the motor command (efference copy), leading to an attenuated perception and neural suppression of the sensory input. Here, we investigated whether and how tools can be incorporated into these predictions. We hypothesized that similar predictive mechanisms are used for both hand and tool use actions, but that additional resources are needed to integrate the tool. During fMRI data acquisition, 19 healthy participants used either their right hand or a tool to hold the handle of a movement device. To manipulate the effect of the efference copy, the handle was moved either actively by participants or passively by the movement device. The sensory outcome, consisting of a real-time video of the hand or tool movement shown on a screen, was presented with varying delays (0-417 ms). Participants reported their perception of such delays. The processing of hand and tool movements yielded largely similar results when comparing active against passive conditions: Active movements were in both cases associated with worse delay detection performances. Moreover, during both hand and tool use actions, active movements led to a downregulation of sensory (somatosensory, visual) areas as well as the right cerebellum and right posterior parietal cortex, as assessed by a conjunction analysis. By contrast, an interaction analysis indicated differential processing of active vs. passive movements in hand vs. tool conditions in the left postcentral gyrus, right middle temporal gyrus (MTG), and bilateral caudate nuclei. Our findings provide behavioral and neural support that hand and tool actions share similar mechanisms for sensory predictions. We propose that the MTG and (sensori)motor areas (postcentral gyrus, caudate nuclei) contribute to these predictions by optimizing them to the physics of the end effector (hand or tool). Collectively, these results suggest that the brain dynamically adjusts sensorimotor predictive models to anticipate the dynamics of the end effector, be it a hand or a tool.

Perceiving your hand moving: BOLD suppression in sensory cortices and the role of the cerebellum in the detection of feedback delays.

Sensory consequences of self-generated as opposed to externally generated movements are perceived as less intense and lead to less neural activity in corresponding sensory cortices, presumably due to predictive mechanisms. Self-generated sensory inputs have been mostly studied in a single modality, using abstract feedback, with control conditions not differentiating efferent from reafferent feedback. Here we investigated the neural processing of (a) naturalistic action-feedback associations of (b) self-generated versus externally generated movements, and (c) how an additional (auditory) modality influences neural processing and detection of delays. Participants executed wrist movements using a passive movement device (PMD) as they watched their movements in real time or with variable delays (0-417 ms). The task was to judge whether there was a delay between the movement and its visual feedback. In the externally generated condition, movements were induced by the PMD to disentangle efferent from reafferent feedback. Half of the trials involved auditory beeps coupled to the onset of the visual feedback. We found reduced BOLD activity in visual, auditory, and somatosensory areas during self-generated compared with externally generated movements in unimodal and bimodal conditions. Anterior and posterior cerebellar areas were engaged for trials in which action-feedback delays were detected for self-generated movements. Specifically, the left cerebellar lobule IX was functionally connected with the right superior occipital gyrus. The results indicate efference copy-based predictive mechanisms specific to self-generated movements, leading to BOLD suppression in sensory areas. In addition, our results support the cerebellum's role in the detection of temporal prediction errors during our actions and their consequences.

Distinct Roles for the Cerebellum, Angular Gyrus, and Middle Temporal Gyrus in Action-Feedback Monitoring.

Action-feedback monitoring is essential to ensure meaningful interactions with the external world. This process involves generating efference copy-based sensory predictions and comparing these with the actual action-feedback. As neural correlates of comparator processes, previous fMRI studies have provided heterogeneous results, including the cerebellum, angular and middle temporal gyrus. However, these studies usually comprised only self-generated actions. Therefore, they might have induced not only action-based prediction errors, but also general sensory mismatch errors. Here, we aimed to disentangle these processes using a custom-made fMRI-compatible movement device, generating active and passive hand movements with identical sensory feedback. Online visual feedback of the hand was presented with a variable delay. Participants had to judge whether the feedback was delayed. Activity in the right cerebellum correlated more positively with delay in active than in passive trials. Interestingly, we also observed activation in the angular and middle temporal gyri, but across both active and passive conditions. This suggests that the cerebellum is a comparator area specific to voluntary action, whereas angular and middle temporal gyri seem to detect more general intersensory conflict. Correlations with behavior and cerebellar activity nevertheless suggest involvement of these temporoparietal areas in processing and awareness of temporal discrepancies in action-feedback monitoring.

Links between Gestures and Multisensory Processing: Individual Differences Suggest a Compensation Mechanism.

Speech-associated gestures represent an important communication modality. However, individual differences in the production and perception of gestures are not well understood so far. We hypothesized that the perception of multisensory action consequences might play a crucial role. Verbal communication involves continuous calibration of audio-visual information produced by the speakers. The effective production and perception of gestures supporting this process could depend on the given capacities to perceive multisensory information accurately. We explored the association between the production and perception of gestures and the monitoring of multisensory action consequences in a sample of 31 participants. We applied a recently introduced gesture scale to assess self-reported gesture production and perception in everyday life situations. In the perceptual experiment, we presented unimodal (visual) and bimodal (visual and auditory) sensory outcomes with various delays after a self-initiated (active) or externally generated (passive) button press. Participants had to report whether they detected a delay between the button press and the visual stimulus. We derived psychometric functions for each condition and determined points of subjective equality, reflecting detection thresholds for delays. Results support a robust link between gesture scores and detection thresholds. Individuals with higher detection thresholds (lower performance) reported more frequent gesture production and perception and furthermore profited more from multisensory information in the experimental task. We propose that our findings indicate a compensational function of multisensory processing as a basis for individual differences in both action outcome monitoring and gesture production and perception in everyday life situations.

Voluntary and Involuntary Movements Widen the Window of Subjective Simultaneity.

Forming a coherent percept of an event requires different sensory inputs originating from the event to be bound. Perceiving synchrony aids in binding of these inputs. In two experiments, we investigated how voluntary movements influence the perception of simultaneity, by measuring simultaneity judgments (SJs) for an audiovisual (AV) stimulus pair triggered by a voluntary button press. In Experiment 1, we manipulated contiguity between the action and its consequences by introducing delays between the button press and the AV stimulus pair. We found a widened window of subjective simultaneity (WSS) when the action-feedback relationship was time contiguous. Introducing a delay narrowed the WSS, suggesting that the wider WSS around the time of an action might facilitate perception of simultaneity. In Experiment 2, we introduced an involuntary condition using an externally controlled button to assess the influence of action-related predictive processes on SJs. We found a widened WSS around the action time, regardless of movement type, supporting the influence of causal relations in the perception of synchrony. Interestingly, the slopes of the psychometric functions in the voluntary condition were significantly steeper than the slopes in the involuntary condition, suggesting a role of action-related predictive mechanisms in making SJs more precise.

Hemispheric differences in the processing of visual consequences of active vs. passive movements: a transcranial direct current stimulation study.

Perceiving the sensory consequences of one's own actions is essential to successfully interact with the environment. Previous studies compared self- (active) and externally generated (passive) movements to investigate the processing of voluntary action-outcomes. Increased temporal binding (intentional binding) as well as increased detection of delays between action and outcome have been observed for active compared to passive movements. Using transcranial direct stimulation (tDCS) it has been shown that left hemispheric anodal stimulation decreased the intentional binding effect. However, whether the left hemisphere contributes to delay detection performance between action and outcome is unknown. We investigated polarization-dependent effects of left and right frontoparietal tDCS on detecting temporal action-outcome discrepancies. We applied anodal and cathodal stimulation to frontal (F3/F4), parietal (CP3/CP4) and frontoparietal (F3/CP4) areas. After stimulation, participants were presented with visual feedback with various delays after a key press. They had to report whether they detected a delay between the key press and the feedback. In half of the trials the key press was self-initiated, in the other half it was externally generated. A main effect of electrode location indicated highest detection performance after frontal stimulation. Furthermore, we found that the advantage for active versus passive conditions was larger for left hemispheric anodal stimulation as compared to cathodal stimulation. Whereas the frontal cortex is related to delay detection performance in general, hemispheric differences seem to support the differentiation of self-initiated versus externally generated movement consequences.

The angular gyrus is a supramodal comparator area in action-outcome monitoring.

Predicting and processing the sensory consequences of one's own actions is essential to enable successful interactions with the environment. Previous studies have suggested that the angular gyrus detects discrepancies between predicted and actual action consequences, at least for unimodal feedback. However, most actions lead to multisensory consequences, raising the question whether previous models can sufficiently explain action-outcome processing. Here, we investigated neural comparator processes during detection of delays between action and unimodal or bimodal consequences in human subjects with fMRI, using parametric and connectivity analyses. Participants had to perform button presses, which led to the presentation of either a dot on the screen, a tone, or both, presented with a variable delay after the button press. Participants were asked to judge whether there was a delay between action and feedback. Activity in the angular gyrus correlated positively with delay for both visual, auditory, and audio-visual action consequences. Furthermore, the angular gyrus was functionally connected with midline structures such as the posterior cingulate cortex and precuneus in all conditions. Our results show that the angular gyrus is (1) a supramodal area, sensitive to delays in multiple modalities, and (2) functionally connected with self-referential areas during delay detection of both unimodal and bimodal action consequences. Overall, our results suggest that the angular gyrus functions as a mediator between perception and interpretation, and that this process is remarkably similar for unimodal and bimodal action consequences.

Predicting the Multisensory Consequences of One's Own Action: BOLD Suppression in Auditory and Visual Cortices.

Predictive mechanisms are essential to successfully interact with the environment and to compensate for delays in the transmission of neural signals. However, whether and how we predict multisensory action outcomes remains largely unknown. Here we investigated the existence of multisensory predictive mechanisms in a context where actions have outcomes in different modalities. During fMRI data acquisition auditory, visual and auditory-visual stimuli were presented in active and passive conditions. In the active condition, a self-initiated button press elicited the stimuli with variable short delays (0-417ms) between action and outcome, and participants had to detect the presence of a delay for auditory or visual outcome (task modality). In the passive condition, stimuli appeared automatically, and participants had to detect the number of stimulus modalities (unimodal/bimodal). For action consequences compared to identical but unpredictable control stimuli we observed suppression of the blood oxygen level depended (BOLD) response in a broad network including bilateral auditory and visual cortices. This effect was independent of task modality or stimulus modality and strongest for trials where no delay was detected (undetected

fMRI-based decoding of reward effects in binocular rivalry.

Binocular rivalry is a phenomenon where the simultaneous presentation of two different stimuli to the two eyes leads to alternating perception of the two stimuli. The temporary dominance of one stimulus over the other is influenced by several factors. Here, we studied the influence of reward on binocular rivalry dynamics and its neural representation in visual cortex. Orthogonal rotating grating stimuli were shown continuously, while monetary reward was given during the conscious perception of one stimulus but not the other. Periods of perceptual dominance were assessed both through participants' subjective report and objectively using functional magnetic resonance imaging and multi-voxel pattern analysis. Results did not confirm previous evidence for an effect of reward on perceptual dominance durations. Exploratory post-hoc analyses indicated that knowledge regarding both the reward contingency and the subjective nature of perceptual alternations may have interfered with potential reward effects on perceptual phase durations, suggesting a moderating role of meta-cognitive awareness in reward-based perceptual inference. Future studies of top-down influences on bistable perception should carefully consider the methodological challenges related to meta-cognitive awareness.

Predicting the sensory consequences of one's own action: First evidence for multisensory facilitation.

Predicting the sensory consequences of our own actions contributes to efficient sensory processing and might help distinguish the consequences of self- versus externally generated actions. Previous research using unimodal stimuli has provided evidence for the existence of a forward model, which explains how such sensory predictions are generated and used to guide behavior. However, whether and how we predict multisensory action outcomes remains largely unknown. Here, we investigated this question in two behavioral experiments. In Experiment 1, we presented unimodal (visual or auditory) and bimodal (visual and auditory) sensory feedback with various delays after a self-initiated buttonpress. Participants had to report whether they detected a delay between their buttonpress and the stimulus in the predefined task modality. In Experiment 2, the sensory feedback and task were the same as in Experiment 1, but in half of the trials the action was externally generated. We observed enhanced delay detection for bimodal relative to unimodal trials, with better performance in general for actively generated actions. Furthermore, in the active condition, the bimodal advantage was largest when the stimulus in the task-irrelevant modality was not delayed-that is, when it was time-contiguous with the action-as compared to when both the task-relevant and task-irrelevant modalities were delayed. This specific enhancement for trials with a nondelayed task-irrelevant modality was absent in the passive condition. These results suggest that a forward model creates predictions for multiple modalities, and consequently contributes to multisensory interactions in the context of action.

Decoding pattern motion information in V1.

When two gratings drifting in different directions are superimposed, the resulting stimulus can be perceived as two overlapping component gratings moving in different directions or as a single pattern moving in one direction. Whilst the motion direction of component gratings is already represented in visual area V1, the majority of previous studies have found processing of pattern motion direction only from visual area V2 onwards. Here, we question these findings using multi-voxel pattern analysis (MVPA). In Experiment 1, we presented superimposed sinusoidal gratings with varying angles between the two component motions. These stimuli were perceived as patterns moving in one of two possible directions. We found that linear support vector machines (SVMs) could generalise across stimuli composed of different component motions to successfully discriminate pattern motion direction from brain activity in V1, V3A and hMT+/V5. This demonstrates the representation of pattern motion information present in these visual areas. This conclusion was verified in Experiment 2, where we manipulated similar plaid stimuli to induce the perception of either a single moving pattern or two separate component gratings. While a classifier could again generalise across stimuli composed of different component motions when they were perceived as a single moving pattern, its performance dropped substantially in the case where components were perceived. Our results indicate that pattern motion direction information is present in V1.

Tactile and visual motion direction processing in hMT+/V5.

The human motion complex hMT+/V5 is activated not only by visual motion, but also by tactile and auditory motion. Whilst direction-selectivity has been found within this complex for visual and auditory stimuli, it is unknown whether hMT+/V5 also contains direction-specific information from the tactile modality. In the current study, we sought to investigate whether hMT+/V5 contains direction-specific information about visual/tactile moving stimuli. Leftward and rightward moving stimuli were presented in the visual and tactile modalities in an event-related fMRI design. Using region-of-interest-based multivariate pattern analysis we could decode the two motion directions for both tactile and visual stimuli in hMT+/V5. The activity patterns of the two modalities differed significantly, indicating that motion direction information from different modalities may be carried by distinct sets of neuronal populations. Our findings show that hMT+/V5 contains specific information about the direction of a moving stimulus in both the tactile and visual modalities, supporting the theory of hMT+/V5 being a multimodal motion area.