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Diedrichsen, J.

Publications and source records attributed to Diedrichsen, J..

At least 55 records · Page 3Linked to original sources

Evaluating brain parcellations using the distance controlled boundary coefficient

One important approach to human brain mapping is to define a set of distinct regions that can be linked to unique functions. Numerous brain parcellations have been proposed, using cytoarchitectonic, structural or functional Magnetic Resonance Imaging (fMRI) data. The intrinsic smoothness of brain data, however, poses a problem for current methods seeking to compare different parcellations. For example, criteria that simply compare within-parcel to between-parcel similarity provide even random parcellations with a high value. Furthermore, the evaluation is biased by the spatial scale of the parcellation. To address this problem, we propose the Distance Controlled Boundary Coefficient (DCBC), an unbiased criterion to evaluate discrete parcellations. We employ this new criterion to evaluate existing parcellations of the human neocortex in their power to predict functional boundaries for an fMRI data set with many different tasks, as well as for resting-state data. We find that common anatomical parcellations do not perform better than chance, suggesting that task-based functional boundaries do not align well with sulcal landmarks. Parcellations based on resting-state fMRI data perform well; in some cases, as well as a parcellation defined on the evaluation data itself. Finally, multi-modal parcellations that combine functional and anatomical criteria perform substantially worse than those based on functional data alone, indicating that functionally homogeneous regions often span major anatomical landmarks. Overall, the DCBC advances the field of functional brain mapping by providing an unbiased metric that compares the predictive ability of different brain parcellations to define brain regions that are functionally maximally distinct.

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Motor planning brings human primary somatosensory cortex into movement-specific preparatory states

Motor planning plays a critical role in producing fast and accurate movement. Yet, the neural processes that occur in human primary motor and somatosensory cortex during planning, and how they relate to those during movement execution, remain poorly understood. Here we used 7T functional magnetic resonance imaging (fMRI) and a delayed movement paradigm to study single finger movement planning and execution. The inclusion of no-go trials and variable delays allowed us to separate what are typically overlapping planning and execution brain responses. Although our univariate results show widespread deactivation during finger planning, multivariate pattern analysis revealed finger-specific activity patterns in contralateral primary somatosensory cortex (S1), which predicted the planned finger action. Surprisingly, these activity patterns were as informative as those found in contralateral primary motor cortex (M1). Control analyses ruled out the possibility that the detected information was an artifact of subthreshold movements during the preparatory delay. Furthermore, we observed that finger-specific activity patterns during planning were highly correlated to those during execution. These findings reveal that motor planning activates the specific S1 and M1 circuits that are engaged during the execution of a finger press, while activity in both regions is overall suppressed. We propose that preparatory states in S1 may improve movement control through changes in sensory processing or via direct influence of spinal motor neurons. Significance statementMotor planning is important for good behavioral performance, yet it is unclear which neural processes underlie the preparation of the nervous system for an upcoming movement. Using high-resolution functional neuroimaging, we investigated how motor planning for finger presses changes the activity state in primary motor and primary somatosensory cortex, and how brain responses during planning and execution relate to each other. We show that planning leads to finger-specific activation in both M1 and S1, which is highly similar to the finger-specific activity patterns elicited during execution. Our findings suggest that S1 is being specifically prepared for an upcoming action, either to actively contribute to the outflowing motor command or to enable action-specific sensory gating.

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Malleability of the cortical hand map following a finger nerve block

Individual fingers in the primary somatosensory cortex (S1) are known to be represented separately and adjacently, forming a cortical hand map. Electrophysiological studies in monkeys show that finger amputation triggers increased selectivity to the neighbouring fingers within the deprived S1, causing local reorganisation. Neuroimaging research in humans, however, shows persistent S1 finger representation of the missing hand, even decades after amputation. We aimed to resolve these apparently contrasting evidence by examining finger representation in humans following pharmacological amputation using single-finger nerve block and 7T neuroimaging. We hypothesised that beneath the apparent selectivity of individual fingers in the hand map, peripheral and central processing is distributed across fingers. If each finger contributes to the cortical representation of the others, then localised input loss will weaken finger representation across the hand map. For the same reason, the non-blocked fingers will stabilise the blocked fingers representation, resulting in persistent representation of the blocked finger. Using univariate selectivity profiling, we replicated the electrophysiological findings of local S1 reorganisation. However, more comprehensive analyses confirmed that local blocking reduced representation of all fingers across the entire hand area. Importantly, multivariate analysis demonstrated that despite input loss, representation of the blocked finger remained persistent and distinct from the unblocked fingers. Computational modelling suggested that the observed findings are driven by distributed processing underlying the topographic map, combined with homeostatic mechanisms. Our findings suggest that the long-standing depiction of the somatosensory hand map is misleading. As such, accounts for map reorganisation, e.g. following amputation, need to be reconsidered.

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Neural correlates of hand-object congruency effects during action planning

Selecting hand actions to manipulate an object is affected both by perceptual factors and by action goals. Affordances are associated with the automatic potentiation of motor representations to an object, independent of the goal of the actor. In previous studies, we have demonstrated an influence of the congruency between hand and object orientations on response times when reaching to turn an object, such as a cup. In this study, we investigated how the representation of hand postures triggered by planning to turn a cup were influenced by this congruency effect, in an fMRI scanning environment. Healthy participants were asked to reach and turn a real cup that was placed in front of them either in an upright orientation or upside down. They were instructed to use a hand orientation that was either congruent or incongruent with the cup orientation. As expected, the motor responses were faster when the hand and cup orientations were congruent. There was increased activity in a network of brain regions involving object-directed actions during action planning, which included bilateral primary and extrastriate visual, medial and superior temporal areas, as well as superior parietal, primary motor and premotor areas in the left hemisphere. Specific activation of the dorsal premotor cortex (PMd) was associated with hand-object orientation congruency during planning, and prior to any action taking place. Activity in that area and its connectivity with the lateral occipito-temporal cortex (LOTC) increased when planning incongruent actions. The increased activity in premotor areas in trials where the orientation of the hand was incongruent to that of the object suggests a role in eliciting competing representations specified by hand postures in LOTC.

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The role of primary motor cortex in sequence learning: resolving conflicting fMRI evidence from repetition suppression and pattern analysis

How does the brain change during learning? In functional magnetic resonance imaging studies, both multivariate pattern analysis and repetition suppression (RS) have been used to detect changes in neuronal representations. In the context of motor sequence learning, the two techniques have provided discrepant findings: pattern analysis showed that only premotor and parietal regions, but not primary motor cortex (M1), develop a representation of trained sequences. In contrast, RS suggested trained sequence representations in all these regions. Here we applied both analysis techniques to a 5-week finger sequence training study, in which participants executed each sequence twice before switching to a different sequence. Both RS and pattern analysis indicated learning-related changes for parietal areas, but only RS showed a difference between trained and untrained sequences in M1. A more fine-grained analysis, however, revealed that the RS effect in M1 reflects a fundamentally different process than in parietal areas. On the first execution, M1 represents especially the first finger of each sequence, likely reflecting preparatory processes. This effect dramatically reduces during the second execution. In contrast, parietal areas represent the identity of a sequence, and this representation stays relatively stable on the second execution. These results suggest that the RS effect does not reflect a trained sequence representation in M1, but rather a preparatory signal for movement initiation. More generally, our study demonstrates that across regions RS can reflect different representational changes in the neuronal population code, emphasizing the importance of combining pattern analysis and RS techniques. Significance statementPrevious studies using pattern analysis have suggested that primary motor cortex (M1) does not represent learnt sequential actions. However, a study using repetition suppression (RS) has reported M1 changes during motor sequence learning. Combining both techniques, we first replicate the discrepancy between them - with learning-related changes in M1 in RS, but not pattern dissimilarities. We further analysed the representational changes with repetition, and found that the RS effects differ across regions. M1s activity represents the starting finger of the sequence, an effect that vanishes with repetition. In contrast, activity patterns in parietal areas exhibit sequence dependency, which persists with repetition. These results demonstrate the importance of combining RS and pattern analysis to understand the function of brain regions.

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The planning horizon for movement sequences

When performing a long chain of actions in rapid sequence, future movements need to be planned concurrently with ongoing action. However, how far ahead we plan, and whether this ability improves with practice, is currently unknown. Here we designed an experiment in which healthy volunteers produced sequences of 14 finger presses quickly and accurately on a keyboard in response to numerical stimuli. On every trial, participants were only shown a fixed number of stimuli ahead of the current keypress. The size of this viewing window varied between 1 (next digit revealed with the pressing of the current key) and 14 (full view of the sequence). Participants practiced the task for five days and their performance was continuously assessed on random sequences. Our results indicate that participants used the available visual information to plan multiple actions into the future, but that the planning horizon was limited: receiving information about more than 3 movements ahead did not result in faster sequence production. Over the course of practice, we found larger performance improvements for larger viewing windows and an expansion of the planning horizon. These findings suggest that the ability to plan future responses during ongoing movement constitutes an important aspect of skillful movement. Based on the results, we propose a framework to investigate the neuronal processes underlying simultaneous planning and execution. Significance StatementAlthough skill learning has typically focused on the training of specific movement sequences, practice improves performance even for random sequences. Here we hypothesize that a fundamental aspect of skilled sequential behavior is the ability to plan multiple actions into the future, both before and during execution. By controlling the amount of visual information available for motor planning, we show that people plan at least three movements beyond current action and that this planning horizon expands with practice. Our findings suggest that coordinating ongoing movement and planning of future actions is an essential component of skilled sequential behavior and offer testable predictions for the neural implementation of online motor planning.

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Structure of population activity in primary motor cortex for single finger flexion and extension

How is the primary motor cortex (M1) organized to control fine finger movements? We investigated the population activity in M1 for single finger flexion and extension, using 7T functional magnetic resonance imaging (fMRI) in female and male human participants, and compared these results to the neural spiking patterns recorded in two male monkeys performing the identical task. fMRI activity patterns were distinct for movements of different fingers, but quite similar for flexion and extension of the same finger. In contrast, spiking patterns in monkeys were quite distinct for both fingers and directions, similar to what was found for muscular activity patterns. The discrepancy between fMRI and electrophysiological measurements can be explained by two (non-mutually exclusive) characteristics of the organization of finger flexion and extension movements. Given that fMRI reflects predominantly input and recurrent activity, the results can be explained by an architecture in which neural populations that control flexion or extension of the same finger produce distinct outputs, but interact tightly with each other and receive similar inputs. Additionally, neurons tuned to different movement directions for the same finger (or combination of fingers) may cluster closely together, while neurons that control different finger combinations may be more spatially separated. When measuring this organization with fMRI at a coarse spatial scale, the activity patterns for flexion and extension of the same finger would appear very similar. Overall, we suggest that the discrepancy between fMRI and electrophysiological measurements provides new insights into the general organization of fine finger movements in M1. Significance statementThe primary motor cortex (M1) is important for producing individuated finger movements. Recent evidence shows that movements that commonly co-occur are associated with more similar activity patterns in M1. Flexion and extension of the same finger, which never co-occur, should therefore be associated with distinct representations. However, using carefully controlled experiments and multivariate analyses, we demonstrate that human fMRI activity patterns for flexion or extension of the same finger are highly similar. In contrast, spiking patterns measured in monkey M1 are clearly distinct. This suggests that populations controlling opposite movements of the same finger, while producing distinct outputs, may cluster together and share inputs and local processing. These results provide testable hypotheses about the organization of hand control in M1.

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fMRI signatures of motor sequence learning: A critical re-evaluation

Despite numerous studies, there is little agreement about what brain changes accompany motor sequence learning, partly because of a general publication bias that favors novel results. We therefore decided to systematically reinvestigate proposed functional magnetic resonance imaging correlates of motor learning in a preregistered longitudinal study with four scanning sessions over 5 weeks of training. Activation decreased more for trained than untrained sequences in premotor and parietal areas, without any evidence of learning-related activation increases. Premotor and parietal regions also exhibited changes in the fine-grained, sequence-specific activation patterns early in learning, which stabilized later. No changes were observed in the primary motor cortex (M1). Overall, our study provides evidence that human motor sequence learning occurs outside of M1. Furthermore, it shows that we cannot expect to find activity increases as an indicator for learning, making subtle changes in activity patterns across weeks the most promising fMRI correlate of training-induced plasticity.

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Repetita iuvant: repetition facilitates online planning of sequential movements

Beyond being essential for long-term motor-skill development, movement repetition has immediate benefits on performance, increasing speed and accuracy of a second execution. While repetition effects have been reported for single reaching movements, it has yet to be determined whether they also occur for movement sequences, and what aspects of sequence production are improved. We addressed these questions in two behavioral experiments using a discrete sequence production (DSP) task in which human volunteers had to perform short sequences of finger movements. In Experiment 1, we presented participants with randomly varying sequences and manipulated 1) whether the same sequence was repeated on successive trials, and 2) whether participants had to execute the sequence (Go), or not (No-Go). We establish that sequence repetition led to immediate improvements in speed without associated accuracy costs. The largest benefit was observed in the middle part of a sequence, suggesting that sequence repetition facilitated online planning. This claim was further supported by Experiment 2, in which we kept a set of sequences fixed throughout the experiment, thus allowing participants to develop sequence-specific learning: once the need for online planning decreased, the benefit of repetition disappeared. Finally, we found that repetition-related improvements only occurred for the trials that had been preceded by sequence production, suggesting that action selection and sequence pre-planning may not be sufficient to reap the benefits of repetition. Together, these results show that repetition can enhance representations at the level of movement sequences (rather than of individual movements) and facilitate online planning. New & NoteworthyEven for overlearned motor skills such as reaching, movement repetition improves performance. How brain processes associated with motor planning or execution benefit from repetition, however, remains unclear. Here we report the novel finding of repetition effects for sequential movements. Our results show that repetition benefits are tied to improved online planning of upcoming sequence elements. We also highlight how actual movement experience appears to be more beneficial than mental rehearsal for observing short-term repetition effects.

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Sensory information from a slipping object elicits a rapid and automatic shoulder response

Humans have the remarkable ability to hold, grasp, and manipulate objects. Previous work has reported rapid and coordinated reactions in hand and shoulder muscles in response to external perturbations to the arm during object manipulation; however, little is known about how somatosensory feedback of an object slipping in the hand influences responses of the arm. We built a hand-held device to stimulate the sensation of slipping at all five fingertips. The device was integrated into an exoskeleton robot that supported it against gravity. The setup allowed us to decouple somatosensory stimulation in the fingers from forces applied to the arm-- two variables that are highly interdependent in real-world scenarios. Fourteen participants performed three experiments in which we measured their arm feedback responses during slip stimulation. Slip stimulations were applied horizontally, in one of two directions, and participants were either instructed to follow the slip direction, or to move the arm in the opposite direction. Participants showed responses within [~]67 ms of slip onset when following the direction of slip, but significantly slower responses when instructed to move in the opposite direction. Arm responses were modulated by the speed but not the distance of the slip. Finally, when slip stimulation was combined with mechanical perturbations to the arm, we found that sensory information from the fingertips significantly modulated the shoulder feedback response. Overall, the results demonstrate the existence of a rapid feedback system that stabilizes hand-held objects.\n\nNEW & NOTHEWORTHYWe tested whether the sensation of an object slipping from the fingers modulates shoulder feedback responses. We found rapid shoulder feedback responses when participants were instructed to follow the slip direction with the arm. Shoulder responses following mechanical joint perturbations were also potentiated when combined with slipping. These results demonstrate the existence of fast and automatic feedback responses in the arm in reaction to sensory input to the fingertips that maintain grip on hand-held objects.

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No evidence for motor recovery-related cortical reorganization after stroke using resting-state fMRI

Cortical reorganization has been suggested as mechanism for recovery after stroke. It has been proposed that a form of cortical reorganization (changes in functional connectivity between brain areas) can be assessed with resting-state fMRI. Here we report the largest longitudinal data-set in terms of overall sessions in 19 patients with subcortical stroke and 11 controls. Patients were imaged up to 5 times over one year. We found no evidence for post-stroke cortical reorganization despite substantial behavioral recovery. These results could be construed as questioning the value of resting-state imaging. Here we argue instead that they are consistent with other emerging reasons to challenge the idea of motor recovery-related cortical reorganization post-stroke when conceived as changes in connectivity between cortical areas.

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Evidence for shared conceptual representations for sign and speech

Do different languages evoke different conceptual representations? If so, greatest divergence might be expected between languages that differ most in structure, such as sign and speech. Unlike speech bilinguals, hearing sign-speech bilinguals use languages conveyed in different modalities. We used functional magnetic resonance imaging and representational similarity analysis (RSA) to quantify the similarity of semantic representations elicited by the same concepts presented in spoken British English and British Sign Language in hearing, early sign-speech bilinguals. We found shared representations for semantic categories in left posterior middle and inferior temporal cortex. Despite shared category representations, the same spoken words and signs did not elicit similar neural patterns. Thus, contrary to previous univariate activation-based analyses of speech and sign perception, we show that semantic representations evoked by speech and sign are only partially shared. This demonstrates the unique perspective that sign languages and RSA provide in understanding how language influences conceptual representation.

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Sequence learning is driven by improvements in motor planning

The ability to perform complex sequences of movements quickly and accurately is critical for many motor skills. While training improves performance in a large variety of motor-sequence tasks, the precise mechanisms behind such improvements are poorly understood. Here we investigated the contribution of single-action selection, sequence pre-planning, online planning, and motor execution to performance in a discrete sequence production (DSP) task. Five visually-presented numbers cued a sequence of five finger presses, which had to be executed as quickly and accurately as possible. To study how sequence planning influenced sequence production, we manipulated the amount of time that participants were given to prepare each sequence by using a forced-response paradigm. Over 4 days, participants were trained on 10 sequences and tested on 80 novel sequences. Our results revealed that participants became faster in selecting individual finger presses. They also preplanned 3-4 sequence items into the future, and the speed of pre-planning improved for trained, but not for untrained, sequences. Because pre-planning capacity remained limited, the remaining sequence elements had to be planned online during sequence execution, a process that also improved with sequence-specific training. Overall, our results support the view that motor sequence learning effects are best characterized by improvements in planning processes that occur both before and concurrently with motor execution.\n\nNew & NoteworthyComplex skills often require the production of sequential movements. While practice improves performance, it remains unclear how these improvements are achieved. Our findings show that learning effects in a sequence production task can be attributed to an enhanced ability to plan upcoming movements. These results shed new light on planning processes in the context of movement sequences, and have important implications for our understanding of the neural mechanisms that underlie skill acquisition.

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Motor skill learning leads to the increase of planning horizon

We investigated motor skill learning using a path tracking task, where human subjects had to track various curved paths at a constant speed while maintaining the cursor within the path width. Subjects accuracy increased with practice, even when tracking novel untrained paths. Using a "searchlight" paradigm, where only a short segment of the path ahead of the cursor was shown, we found that subjects with a higher tracking skill differed from the novice subjects in two respects. First, they had lower motor variability, in agreement with previous findings. Second, they took a longer section of the future path into account when performing the task, i.e. had a longer planning horizon. We estimate that between one third and one half of the performance increase was due to the increase in planning horizon. An optimal control model with a fixed horizon (receding horizon control) that increases with tracking skill quantitatively captured the subjects movement behaviour. These findings demonstrate that human subjects not only increase their motor acuity but also their planning horizon when acquiring a motor skill. New and NoteworthyWe show that when learning a motor skill humans are using information about the environment from an increasingly longer amount of the movement path ahead to improve performance. Crucial features of the behavioural performance can be captured by modelling the behavioural data with a receding horizon optimal control model.

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A Multi-Domain Task Battery Reveals Functional Boundaries in the Human Cerebellum

There is compelling evidence that the human cerebellum is engaged in a wide array of motor and cognitive tasks. A fundamental question centers on whether the cerebellum is organized into distinct functional sub-regions. To address this question, we employed a rich task battery, designed to tap into a broad range of cognitive processes. During four functional magnetic resonance imaging (fMRI) sessions, participants performed a battery of 26 diverse tasks comprising 47 unique conditions. Using the data from this multi-domain task battery, we derived a comprehensive functional parcellation of the cerebellar cortex, and evaluated it by predicting functional boundaries in a novel set of tasks. The new parcellation successfully identified distinct functional sub-regions, providing significant improvements over existing parcellations derived from task-free data. Lobular boundaries, commonly used to summarize functional data, did not coincide with functional subdivisions. This multi-domain task approach offers novel insights into the functional heterogeneity of the cerebellar cortex.

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Parcellation of motor sequence representations in the human neocortex

While previous studies have revealed an extended network of cortical regions associated with motor sequence production, the specific role of each of these areas is still elusive. To address this issue, we designed a novel behavioural paradigm that allowed us to experimentally manipulate the structure of motor sequences representations in individual participants. We then conducted fMRI while participants executed 8 trained sequences to examine how this structure is reflected in the associated activity patterns. Both model-based and model-free approaches revealed a clear distinction between primary and non-primary motor cortices in their representational contents, with M1 specifically representing individual finger movements, and premotor and parietal cortices showing a mixture of chunk, sequence and finger transition representations. Using model-free representational parcellation, we could divide these non-primary motor cortices into separate clusters, each with a unique representation along the stimulus-to-action gradient. These results provide new insights into how human neocortex organizes movement sequences.

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The effects of habits on motor skill learning

Skill learning involves the formation of stable motor patterns. In musical and athletic training, however, these stable motor habits can also impede the attainment of higher levels of performance. We developed an experimental paradigm to induce a specific motor pattern in the context of a discrete sequence production task and to investigate how these habits affect performance over a 3-week training period. Participants initially practiced small segments of 2 to 3 finger movements ("chunks") and then learned longer sequences composed of these chunks. This initial training induced a persistent temporal pattern during execution, with shorter inter-press-intervals within a chunk and longer ones at chunk boundaries. This pattern remained stable during the subsequent 10 days of training, in which participants were asked to produce the sequence as fast as possible from memory. The habit was also preserved when the sequences were directly displayed, removing the need for memory recall. We were able to induce chunking patterns that were either beneficial or detrimental to performance by taking into consideration the biomechanical constraints of the sequences. While we observed an overall reduction in the detrimental effect of the disadvantageous chunking instructions with training, our results show that the degree to which these detrimental chunk structures were maintained, was predictive of lower levels of final performance. In sum, we were able to induce beneficial and detrimental motor habits in a motor sequence production task and show that these initial instructions influenced performance outcomes over a prolonged period of time. Significance StatementA habit is defined as an automatized action that resists modification once sufficiently established. Preventing bad habits, while reinforcing good habits, is a key objective when teaching new motor skills. While habit formation is an integral part of motor skill acquisition, previous research has focused on habit formation in terms of action selection. In this paper, we examine habit formation in terms of motor skill execution, after the action has been selected. We were able to induce beneficial or detrimental motor habits in the production of motor sequences. Habits were stable over a prolonged training period. Our results demonstrate how cognitive instruction can lead to persistent motor habits and we explore how these habits are potentially modified with training.

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Ipsilateral finger representations are engaged in active movement, but not sensory processing

Hand and finger movements are mostly controlled through crossed corticospinal projections from the contralateral hemisphere. During unimanual movements, activity in the contralateral hemisphere is increased while the ipsilateral hemisphere is suppressed below resting baseline. Despite this suppression, unimanual movements can be decoded from ipsilateral activity alone. This indicates that ipsilateral activity patterns represent parameters of ongoing movement, but the origin and functional relevance of these representations is unclear. Here, we asked whether human ipsilateral representations are caused by active movement, or whether they are driven by sensory input. Participants alternated between performing single finger presses and having fingers passively stimulated, while we recorded brain activity using high-field (7T) functional imaging. We contrasted active and passive finger representations in sensorimotor areas of ipsilateral and contralateral hemispheres. Finger representations in the contralateral hemisphere were equally strong under passive and active conditions, highlighting the importance of sensory information in feedback control. In contrast, ipsilateral finger representations were stronger during active presses. Furthermore, the spatial distribution of finger representations differed between hemispheres: the contralateral hemisphere showed the strongest finger representations in Brodmann area 3a and 3b, while the ipsilateral hemisphere exhibited stronger representations in premotor and parietal areas. This suggests that finger representations in the two hemispheres have different origins - contralateral representations are driven by both active movement and sensory stimulation, whereas ipsilateral representations are mainly engaged during active movement. This suggests that a possible contribution of the ipsilateral hemisphere lies in movement planning, rather than in the dexterous feedback control of the movement.\n\nSignificance statementMovements of the human body are mostly controlled by contralateral cortical regions. However, activity in ipsilateral sensorimotor regions is also modulated during active movements. The origin and functional relevance of these ipsilateral representations is unclear. Here we used high-field neuroimaging to investigate how human contralateral and ipsilateral hemispheres represent active finger presses and passive finger stimulation. We report that while the contralateral hemisphere was equally strongly recruited during active and passive conditions, the ipsilateral hemisphere was mostly recruited during active movement. We propose that the ipsilateral hemisphere may play a role in bilateral movement planning.

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