Perception of spatial orientation is thought to rely on the brain's integration of visual, vestib... more Perception of spatial orientation is thought to rely on the brain's integration of visual, vestibular, proprioceptive, and somatosensory signals, as well as internal beliefs. When one of these signals breaks down, such as the vestibular signal in bilateral vestibulopathy, patients start compensating by relying more on the remaining cues. How these signals are reweighted in this integration process is difficult to establish, since they cannot be measured in isolation during natural tasks, are inherently noisy, and can be ambiguous or in conflict. Here, we review our recent work, combining experimental psychophysics with a reverse engineering approach, based on Bayesian inference principles, to quantify sensory noise levels and optimal (re)weighting at the individual subject level, in both patients with bilateral vestibular deficits and healthy controls. We show that these patients reweight the remaining sensory information, relying more on visual and other nonvestibular information than healthy controls in the perception of spatial orientation. This quantification approach could improve diagnostics and prognostics of multisensory integration deficits in vestibular patients, and contribute to an evaluation of rehabilitation therapies directed toward specific training programs.
bioRxiv (Cold Spring Harbor Laboratory), Mar 9, 2023
Voice production can be a whole-body affair: Upper limb movements physically impact the voice in ... more Voice production can be a whole-body affair: Upper limb movements physically impact the voice in steady-state vocalization, speaking, and singing. This is supposedly due to biomechanical impulses on the chest-wall, affecting subglottal pressure. Unveiling such biomechanics is important, as humans gesture with their hands in a synchronized way with speaking. Here we assess biomechanical interactions between arm movements and the voice, by measurement of key (respiratoryrelated) muscles with electromyography (EMG) during different types of upper limb movement while measuring the bodys center of mass. We show that gesture-related muscle activations scale with positive peaks in the voices amplitude. Some of these muscles also strongly associate with changes in the center mass, confirming that gesture-vocal coupling partly arises due to posture-related muscle activity. If replicated, these results suggest an evolutionary ancient gesture-vocal connection at the level of biomechanics. These preliminary results will support a pre-registration of analyses for a larger-scale confirmatory study.
Motor costs influence movement selection. These costs could change when movements are adapted in ... more Motor costs influence movement selection. These costs could change when movements are adapted in response to errors. When the motor system attributes the encountered errors to an external cause, appropriate movement selection requires an update of the movement goal, which prompts the selection of a different control policy. However, when errors are attributed to an internal cause, the initially selected control policy could remain unchanged, but the internal forward model of the body needs to be updated, resulting in an online correction of the movement. We hypothesized that external attribution of errors leads to the selection of a different control policy, and thus to a change in the expected cost of movements. This should also affect subsequent motor decisions. Conversely, internal attribution of errors may (initially) only evoke online corrections, and thus is expected to leave the motor decision process unchanged. We tested this hypothesis using a saccadic adaptation paradigm, designed to change the relative motor cost of two targets. Motor decisions were measured using a target selection task between the two saccadic targets before and after adaptation. Adaptation was induced by either abrupt or gradual perturbation schedules, which are thought to induce more external or internal attribution of errors, respectively. By taking individual variability into account, our results show that saccadic decisions shift towards the least costly target after adaptation, but only when the perturbation is abruptly, and not gradually, introduced. We suggest that credit assignment of errors not only influences motor adaptation but also subsequent motor decisions.
Many sensorimotor activities have a time constraint for successful completion. In this case, any ... more Many sensorimotor activities have a time constraint for successful completion. In this case, any time devoted to sensory processing is at the expense of time available for motor execution. Earlier studies have explored how this competition between sensory processing and motor execution is resolved by using experimental designs that segregate the sensing and acting phase of the task. It was found that participants switch from the sensing to the acting stage such that the overall (sensorimotor) uncertainty in the outcome of the task is minimized. An unexplained observation in these studies is the substantial variability in switching times. We investigated the variability in switching time by correlating it with the underlying sensorimotor uncertainty. To this end, we used a modified version of the virtual ball catching paradigm proposed by Faisal & Wolpert (2009). Subjects were instructed to catch a ball, but as soon as they initiated their movement the ball disappeared. We extended the range of horizontal velocities and used two different start positions of the ball to quantify the dependence of sensory uncertainty on ball velocity. Velocity dependence of sensory uncertainty allowed us to manipulate sensory uncertainty and hence the sensorimotor uncertainty. We found that the variability in switching times is correlated with two factors. Firstly, variability in switching times is greater when variation in switching time has only minimal effects on sensorimotor uncertainty. Secondly, variability in switching times is greater when the sensorimotor uncertainty is larger. Our results suggest that the variability in switching time reflects an uncertainty-driven exploratory process.
bioRxiv (Cold Spring Harbor Laboratory), Jul 13, 2022
Generalization in motor learning refers to the transfer of a learned compensation to other releva... more Generalization in motor learning refers to the transfer of a learned compensation to other relevant contexts. The generalization function is typically assumed to be of Gaussian shape, centered on the planned motion, although more recent studies associate generalization with the actual motion. Because motor learning is thought to involve multiple adaptive processes with different time constants, we hypothesized that these processes have different time-dependent contributions to the generalization. Guided by a model-based approach, the objective of the present study was to experimentally examine these contributions. We first reformulated a validated two-state adaptation model as a combination of weighted motor primitives, each specified as a Gaussian-shaped tuning function. Adaptation in this model is achieved by updating individual weights of the primitives of the fast and slow adaptive process separately. Depending on whether updating occurred in a planreferenced or a motion-referenced manner, the model predicted distinct contributions to the overall generalization by the slow and fast process. We tested 23 participants in a reach adaptation task, using a spontaneous recovery paradigm consisting of five successive blocks of a long adaptation phase to a viscous force field, a short adaptation phase with the opposite force, and an error-clamp phase. Generalization was assessed in eleven movement directions relative to the trained target direction. Results of our participant population fell within a continuum of evidence for plan-referenced to evidence for motion-referenced updating. This mixture may reflect the differential weighting of explicit and implicit compensation strategies among participants.
Full body-motion biases decisions of hand choice. We examined the signatures of this bias in hand... more Full body-motion biases decisions of hand choice. We examined the signatures of this bias in hand preference in corticospinal excitability before a reach target was presented. Our results show that behavior and corticospinal excitability modulate depending on the state of the body in motion. This suggests that information about body motion penetrates deeply within the motor system.
Pointing movements can serve instrumental goals ('pointing to press a button') or communicative g... more Pointing movements can serve instrumental goals ('pointing to press a button') or communicative goals ('pointing to indicate to someone which button to press'). Previous work has shown that communicative pointing follows different trajectories, and has different end points than instrumental pointing movements, depending on the addressee's spatial location. This suggests that motor control processes are affected by communicative intentions, but the nature of this interface remains unknown. Using optimal feedback control theory, we construct a model of instrumental pointing, and explore how this model can be adjusted to reproduce the dependency between communicative trajectories and addressees' locations. Our results show that the variations in end points cannot account for those trajectories. Instead, the kinematic data are best explained by 'perspective taking' on the part of the communicator, i.e., communicative pointing movements seem to be planned in a frame of reference that is adjusted to the addressee's point of view.
Contemporary motor control theories propose competition between multiple motor plans before the w... more Contemporary motor control theories propose competition between multiple motor plans before the winning command is executed. While most competitions are completed before movement onset, movements are often initiated before the competition has been resolved. An example of this is saccadic averaging, wherein the eyes land at an intermediate location between two visual targets. Behavioral and neurophysiological signatures of competing motor commands have also been reported for reaching movements, but debate remains about whether such signatures attest to an unresolved competition, arise from averaging across many trials, or reflect a strategy to optimize behavior given task constraints. Here, we recorded EMG activity from an upper limb muscle (m. pectoralis) while 12 (8 female) participants performed an immediate response reach task, freely choosing between one of two identical and suddenly presented visual targets. On each trial, muscle recruitment showed two distinct phases of directionally tuned activity. In the first wave, time-locked ∼100 ms of target presentation, muscle activity was clearly influenced by the nonchosen target, reflecting a competition between reach commands that was biased in favor of the ultimately chosen target. This resulted in an initial movement intermediate between the two targets. In contrast, the second wave, time-locked to voluntary reach onset, was not biased toward the nonchosen target, showing that the competition between targets was resolved. Instead, this wave of activity compensated for the averaging induced by the first wave. Thus, single-trial analysis reveals an evolution in how the nonchosen target differentially influences the first and second wave of muscle activity.SIGNIFICANCE STATEMENTContemporary theories of motor control suggest that multiple motor plans compete for selection before the winning command is executed. Evidence for this is found in intermediate reach movements toward two potential target locations, but recent findings have challenged this notion by arguing that intermediate reaching movements reflect an optimal response strategy. By examining upper limb muscle recruitment during a free-choice reach task, we show early recruitment of a suboptimal averaged motor command to the two targets that subsequently transitions to a single motor command that compensates for the initially averaged motor command. Recording limb muscle activity permits single-trial resolution of the dynamic influence of the nonchosen target through time.
Estimation of the orientation of the head relative to the earth's vertical is thought to rely on ... more Estimation of the orientation of the head relative to the earth's vertical is thought to rely on the integration of vestibular and visual cues. The role of visual cues can be tested using a rod-and-frame task in which a global visual scene, typically a square frame, is displayed at different orientations together with a rod whose perceived direction is a proxy for the head-in-space estimate. While it is known that the frame biases this percept, and hence the subjective visual vertical, the possible role of the rod itself in this processing has not been examined. Current models about spatial orientation assume that the visual orientation of the rod and its uncertainty play no role in the visual-vestibular integration process, but are only involved in the transformation that yields rod orientation in space, thereby contributing additive noise to the subjective visual vertical. Here we tested the validity of this assumption in the rod-and-frame task by replacing the rod with an ellipse whose orientation uncertainty was manipulated by varying its eccentricity (i.e., making the ellipse more or less rounded). Using a psychophysical approach, subjects performed this ellipse-and-frame task for three different eccentricities of the ellipse (0.74, 0.82, 0.99) and three frame orientations (Ă€17.58, 08, 17.58). Results show that ellipse eccentricity affects the uncertainty but not the bias of the subjective visual vertical, suggesting that the ellipse does not interact with the frame in global visual processing but contributes additive noise in computing its orientation in world coordinates.
ABSTRACTVoice production can be a whole-body affair: Upper limb movements physically impact the v... more ABSTRACTVoice production can be a whole-body affair: Upper limb movements physically impact the voice in steady-state vocalization, speaking, and singing. This is supposedly due to biomechanical impulses on the chest-wall, affecting subglottal pressure. Unveiling such biomechanics is important, as humans gesture with their hands in a synchronized way with speaking. Here we assess biomechanical interactions between arm movements and the voice, by measurement of key (respiratory-related) muscles with electromyography (EMG) during different types of upper limb movement while measuring the bodys center of mass. We show that gesture-related muscle activations scale with positive peaks in the voices amplitude. Some of these muscles also strongly associate with changes in the center mass, confirming that gesture-vocal coupling partly arises due to posture-related muscle activity. If replicated, these results suggest an evolutionary ancient gesture-vocal connection at the level of biomechan...
Within predictive processing two kinds of learning can be distinguished: parameter learning and s... more Within predictive processing two kinds of learning can be distinguished: parameter learning and structure learning. In Bayesian parameter learning, parameters under a specific generative model are continuously being updated in light of new evidence. However, this learning mechanism cannot explain how new parameters are added to a model. Structure learning, unlike parameter learning, makes structural changes to a generative model by altering its causal connections or adding or removing parameters. Whilst these two types of learning have recently been formally differentiated, they have not been empirically distinguished. The aim of this research was to empirically differentiate between parameter learning and structure learning on the basis of how they affect pupil dilation. Participants took part in a within-subject computer-based learning experiment with two phases. In the first phase, participants had to learn the relationship between cues and target stimuli. In the second phase, th...
Notice of republication This article was republished on April 1, 2022, to update the author bylin... more Notice of republication This article was republished on April 1, 2022, to update the author byline and list all members of the group consortium. Please download this article again to view the correct version. The originally published, uncorrected article and the republished, corrected articles are provided here for reference.
The endeavor to understand the human brain has seen more progress in the last few decades than in... more The endeavor to understand the human brain has seen more progress in the last few decades than in the previous two millennia. Still, our understanding of how the human brain relates to behavior in the real world and how this link is modulated by biological, social, and environmental factors is limited. To address this, we designed the Healthy Brain Study (HBS), an interdisciplinary, longitudinal, cohort study based on multidimensional, dynamic assessments in both the laboratory and the real world. Here, we describe the rationale and design of the currently ongoing HBS. The HBS is examining a population-based sample of 1,000 healthy participants (age 30–39) who are thoroughly studied across an entire year. Data are collected through cognitive, affective, behavioral, and physiological testing, neuroimaging, bio-sampling, questionnaires, ecological momentary assessment, and real-world assessments using wearable devices. These data will become an accessible resource for the scientific c...
Within predictive processing learning is construed as Bayesian model updating with the degree of ... more Within predictive processing learning is construed as Bayesian model updating with the degree of certainty for different existing hypotheses changing in light of new evidence. Bayesian model updating, however, cannot explain how new hypotheses are added to a model. Model revision, unlike model updating, makes structural changes to a generative model by altering its causal connections or adding or removing hypotheses. Whilst model updating and model revision have recently been formally differentiated, they have not been empirically distinguished. The aim of this research was to empirically differentiate between model updating and revision on the basis of how they affect prediction errors and predictions over time. To study this, participants took part in a within-subject computer-based learning experiment with two phases: updating and revision. In the updating phase, participants had to predict the relationship between cues and target stimuli and in the revision phase, they had to co...
Behavioral studies have shown that humans account for inertial acceleration in their decisions of... more Behavioral studies have shown that humans account for inertial acceleration in their decisions of hand choice when reaching during body motion. Physiologically, it is unclear at what stage of movement preparation information about body motion is integrated in the process of hand selection. Here, we addressed this question by applying transcranial magnetic stimulation over motor cortex (M1) of human participants who performed a preferential reach task while they were sinusoidally translated on a linear motion platform. If M1 only represents a read-out of the final hand choice, we expect the body motion not to affect the MEP amplitude. If body motion biases the hand selection process prior to target onset, we expect corticospinal excitability to modulate with the phase of the motion, with larger MEP amplitudes for phases that show a bias to using the right hand. Behavioral results replicate our earlier findings of a sinusoidal modulation of hand choice bias with motion phase. MEP ampl...
Contemporary theories of motor control have suggested that multiple motor commands compete for ac... more Contemporary theories of motor control have suggested that multiple motor commands compete for action selection. While most of these competitions are completed prior to movement onset, averaged saccadic eye movements that land at an intermediate location between two visual targets are thought to arise when a movement is initiated prior to the resolution of the competition. In contrast, while averaged reach movements have been reported, there is still debate on whether averaged reach movements are the result of a resolved competition between two potential actions or a strategic behavior that optimally incorporates the current task demands. Here, we use a reach version of the paradigm that has previously shown to elicit saccadic averaging to examine whether similar averaging occurs based on neuromuscular activity of an upper limb muscle. On a single trial basis, we observed a temporal evolution of the two competing motor commands during a free-choice reach task to one of two visual ta...
Recent computational theories and behavioral observations suggest that motor learning is supporte... more Recent computational theories and behavioral observations suggest that motor learning is supported by multiple adaptation processes, operating on different timescales, but direct neural evidence is lacking. We tested this hypothesis by applying transcranial magnetic stimulation over motor cortex in 16 human subjects during a validated reach adaptation task. Motor-evoked potentials (MEPs) and cortical silent periods (CSPs) were recorded from the biceps brachii to assess modulations of corticospinal excitability as indices for corticospinal plasticity. Guided by a two-state adaptation model, we show that the MEP reflects an adaptive process that learns quickly but has poor retention, while the CSP correlates with a process that responds more slowly but retains information well. These results provide a physiological link between models of motor learning and distinct changes in corticospinal excitability. Our findings support the relationship between corticospinal gain modulations and t...
For the brain to decide on a reaching movement, it needs to select which hand to use. A number of... more For the brain to decide on a reaching movement, it needs to select which hand to use. A number of body-centered factors affect this decision, such as the anticipated movement costs of each arm, recent choice success, handedness, and task demands. While the position of each hand relative to the target is also known to be an important spatial factor, it is unclear which reference frames coordinate the spatial aspects in the decisions of hand choice. Here we tested the role of gaze- and head-centered reference frames in a hand selection task. With their head and gaze oriented in different directions, we measured hand choice of 19 right-handed subjects instructed to make unimanual reaching movements to targets at various directions relative to their body. Using an adaptive procedure, we determined the target angle that led to equiprobable right/left hand choices. When gaze remained fixed relative to the body this balanced target angle shifted systematically with head orientation, and wh...
Perception of spatial orientation is thought to rely on the brain's integration of visual, vestib... more Perception of spatial orientation is thought to rely on the brain's integration of visual, vestibular, proprioceptive, and somatosensory signals, as well as internal beliefs. When one of these signals breaks down, such as the vestibular signal in bilateral vestibulopathy, patients start compensating by relying more on the remaining cues. How these signals are reweighted in this integration process is difficult to establish, since they cannot be measured in isolation during natural tasks, are inherently noisy, and can be ambiguous or in conflict. Here, we review our recent work, combining experimental psychophysics with a reverse engineering approach, based on Bayesian inference principles, to quantify sensory noise levels and optimal (re)weighting at the individual subject level, in both patients with bilateral vestibular deficits and healthy controls. We show that these patients reweight the remaining sensory information, relying more on visual and other nonvestibular information than healthy controls in the perception of spatial orientation. This quantification approach could improve diagnostics and prognostics of multisensory integration deficits in vestibular patients, and contribute to an evaluation of rehabilitation therapies directed toward specific training programs.
bioRxiv (Cold Spring Harbor Laboratory), Mar 9, 2023
Voice production can be a whole-body affair: Upper limb movements physically impact the voice in ... more Voice production can be a whole-body affair: Upper limb movements physically impact the voice in steady-state vocalization, speaking, and singing. This is supposedly due to biomechanical impulses on the chest-wall, affecting subglottal pressure. Unveiling such biomechanics is important, as humans gesture with their hands in a synchronized way with speaking. Here we assess biomechanical interactions between arm movements and the voice, by measurement of key (respiratoryrelated) muscles with electromyography (EMG) during different types of upper limb movement while measuring the bodys center of mass. We show that gesture-related muscle activations scale with positive peaks in the voices amplitude. Some of these muscles also strongly associate with changes in the center mass, confirming that gesture-vocal coupling partly arises due to posture-related muscle activity. If replicated, these results suggest an evolutionary ancient gesture-vocal connection at the level of biomechanics. These preliminary results will support a pre-registration of analyses for a larger-scale confirmatory study.
Motor costs influence movement selection. These costs could change when movements are adapted in ... more Motor costs influence movement selection. These costs could change when movements are adapted in response to errors. When the motor system attributes the encountered errors to an external cause, appropriate movement selection requires an update of the movement goal, which prompts the selection of a different control policy. However, when errors are attributed to an internal cause, the initially selected control policy could remain unchanged, but the internal forward model of the body needs to be updated, resulting in an online correction of the movement. We hypothesized that external attribution of errors leads to the selection of a different control policy, and thus to a change in the expected cost of movements. This should also affect subsequent motor decisions. Conversely, internal attribution of errors may (initially) only evoke online corrections, and thus is expected to leave the motor decision process unchanged. We tested this hypothesis using a saccadic adaptation paradigm, designed to change the relative motor cost of two targets. Motor decisions were measured using a target selection task between the two saccadic targets before and after adaptation. Adaptation was induced by either abrupt or gradual perturbation schedules, which are thought to induce more external or internal attribution of errors, respectively. By taking individual variability into account, our results show that saccadic decisions shift towards the least costly target after adaptation, but only when the perturbation is abruptly, and not gradually, introduced. We suggest that credit assignment of errors not only influences motor adaptation but also subsequent motor decisions.
Many sensorimotor activities have a time constraint for successful completion. In this case, any ... more Many sensorimotor activities have a time constraint for successful completion. In this case, any time devoted to sensory processing is at the expense of time available for motor execution. Earlier studies have explored how this competition between sensory processing and motor execution is resolved by using experimental designs that segregate the sensing and acting phase of the task. It was found that participants switch from the sensing to the acting stage such that the overall (sensorimotor) uncertainty in the outcome of the task is minimized. An unexplained observation in these studies is the substantial variability in switching times. We investigated the variability in switching time by correlating it with the underlying sensorimotor uncertainty. To this end, we used a modified version of the virtual ball catching paradigm proposed by Faisal & Wolpert (2009). Subjects were instructed to catch a ball, but as soon as they initiated their movement the ball disappeared. We extended the range of horizontal velocities and used two different start positions of the ball to quantify the dependence of sensory uncertainty on ball velocity. Velocity dependence of sensory uncertainty allowed us to manipulate sensory uncertainty and hence the sensorimotor uncertainty. We found that the variability in switching times is correlated with two factors. Firstly, variability in switching times is greater when variation in switching time has only minimal effects on sensorimotor uncertainty. Secondly, variability in switching times is greater when the sensorimotor uncertainty is larger. Our results suggest that the variability in switching time reflects an uncertainty-driven exploratory process.
bioRxiv (Cold Spring Harbor Laboratory), Jul 13, 2022
Generalization in motor learning refers to the transfer of a learned compensation to other releva... more Generalization in motor learning refers to the transfer of a learned compensation to other relevant contexts. The generalization function is typically assumed to be of Gaussian shape, centered on the planned motion, although more recent studies associate generalization with the actual motion. Because motor learning is thought to involve multiple adaptive processes with different time constants, we hypothesized that these processes have different time-dependent contributions to the generalization. Guided by a model-based approach, the objective of the present study was to experimentally examine these contributions. We first reformulated a validated two-state adaptation model as a combination of weighted motor primitives, each specified as a Gaussian-shaped tuning function. Adaptation in this model is achieved by updating individual weights of the primitives of the fast and slow adaptive process separately. Depending on whether updating occurred in a planreferenced or a motion-referenced manner, the model predicted distinct contributions to the overall generalization by the slow and fast process. We tested 23 participants in a reach adaptation task, using a spontaneous recovery paradigm consisting of five successive blocks of a long adaptation phase to a viscous force field, a short adaptation phase with the opposite force, and an error-clamp phase. Generalization was assessed in eleven movement directions relative to the trained target direction. Results of our participant population fell within a continuum of evidence for plan-referenced to evidence for motion-referenced updating. This mixture may reflect the differential weighting of explicit and implicit compensation strategies among participants.
Full body-motion biases decisions of hand choice. We examined the signatures of this bias in hand... more Full body-motion biases decisions of hand choice. We examined the signatures of this bias in hand preference in corticospinal excitability before a reach target was presented. Our results show that behavior and corticospinal excitability modulate depending on the state of the body in motion. This suggests that information about body motion penetrates deeply within the motor system.
Pointing movements can serve instrumental goals ('pointing to press a button') or communicative g... more Pointing movements can serve instrumental goals ('pointing to press a button') or communicative goals ('pointing to indicate to someone which button to press'). Previous work has shown that communicative pointing follows different trajectories, and has different end points than instrumental pointing movements, depending on the addressee's spatial location. This suggests that motor control processes are affected by communicative intentions, but the nature of this interface remains unknown. Using optimal feedback control theory, we construct a model of instrumental pointing, and explore how this model can be adjusted to reproduce the dependency between communicative trajectories and addressees' locations. Our results show that the variations in end points cannot account for those trajectories. Instead, the kinematic data are best explained by 'perspective taking' on the part of the communicator, i.e., communicative pointing movements seem to be planned in a frame of reference that is adjusted to the addressee's point of view.
Contemporary motor control theories propose competition between multiple motor plans before the w... more Contemporary motor control theories propose competition between multiple motor plans before the winning command is executed. While most competitions are completed before movement onset, movements are often initiated before the competition has been resolved. An example of this is saccadic averaging, wherein the eyes land at an intermediate location between two visual targets. Behavioral and neurophysiological signatures of competing motor commands have also been reported for reaching movements, but debate remains about whether such signatures attest to an unresolved competition, arise from averaging across many trials, or reflect a strategy to optimize behavior given task constraints. Here, we recorded EMG activity from an upper limb muscle (m. pectoralis) while 12 (8 female) participants performed an immediate response reach task, freely choosing between one of two identical and suddenly presented visual targets. On each trial, muscle recruitment showed two distinct phases of directionally tuned activity. In the first wave, time-locked ∼100 ms of target presentation, muscle activity was clearly influenced by the nonchosen target, reflecting a competition between reach commands that was biased in favor of the ultimately chosen target. This resulted in an initial movement intermediate between the two targets. In contrast, the second wave, time-locked to voluntary reach onset, was not biased toward the nonchosen target, showing that the competition between targets was resolved. Instead, this wave of activity compensated for the averaging induced by the first wave. Thus, single-trial analysis reveals an evolution in how the nonchosen target differentially influences the first and second wave of muscle activity.SIGNIFICANCE STATEMENTContemporary theories of motor control suggest that multiple motor plans compete for selection before the winning command is executed. Evidence for this is found in intermediate reach movements toward two potential target locations, but recent findings have challenged this notion by arguing that intermediate reaching movements reflect an optimal response strategy. By examining upper limb muscle recruitment during a free-choice reach task, we show early recruitment of a suboptimal averaged motor command to the two targets that subsequently transitions to a single motor command that compensates for the initially averaged motor command. Recording limb muscle activity permits single-trial resolution of the dynamic influence of the nonchosen target through time.
Estimation of the orientation of the head relative to the earth's vertical is thought to rely on ... more Estimation of the orientation of the head relative to the earth's vertical is thought to rely on the integration of vestibular and visual cues. The role of visual cues can be tested using a rod-and-frame task in which a global visual scene, typically a square frame, is displayed at different orientations together with a rod whose perceived direction is a proxy for the head-in-space estimate. While it is known that the frame biases this percept, and hence the subjective visual vertical, the possible role of the rod itself in this processing has not been examined. Current models about spatial orientation assume that the visual orientation of the rod and its uncertainty play no role in the visual-vestibular integration process, but are only involved in the transformation that yields rod orientation in space, thereby contributing additive noise to the subjective visual vertical. Here we tested the validity of this assumption in the rod-and-frame task by replacing the rod with an ellipse whose orientation uncertainty was manipulated by varying its eccentricity (i.e., making the ellipse more or less rounded). Using a psychophysical approach, subjects performed this ellipse-and-frame task for three different eccentricities of the ellipse (0.74, 0.82, 0.99) and three frame orientations (Ă€17.58, 08, 17.58). Results show that ellipse eccentricity affects the uncertainty but not the bias of the subjective visual vertical, suggesting that the ellipse does not interact with the frame in global visual processing but contributes additive noise in computing its orientation in world coordinates.
ABSTRACTVoice production can be a whole-body affair: Upper limb movements physically impact the v... more ABSTRACTVoice production can be a whole-body affair: Upper limb movements physically impact the voice in steady-state vocalization, speaking, and singing. This is supposedly due to biomechanical impulses on the chest-wall, affecting subglottal pressure. Unveiling such biomechanics is important, as humans gesture with their hands in a synchronized way with speaking. Here we assess biomechanical interactions between arm movements and the voice, by measurement of key (respiratory-related) muscles with electromyography (EMG) during different types of upper limb movement while measuring the bodys center of mass. We show that gesture-related muscle activations scale with positive peaks in the voices amplitude. Some of these muscles also strongly associate with changes in the center mass, confirming that gesture-vocal coupling partly arises due to posture-related muscle activity. If replicated, these results suggest an evolutionary ancient gesture-vocal connection at the level of biomechan...
Within predictive processing two kinds of learning can be distinguished: parameter learning and s... more Within predictive processing two kinds of learning can be distinguished: parameter learning and structure learning. In Bayesian parameter learning, parameters under a specific generative model are continuously being updated in light of new evidence. However, this learning mechanism cannot explain how new parameters are added to a model. Structure learning, unlike parameter learning, makes structural changes to a generative model by altering its causal connections or adding or removing parameters. Whilst these two types of learning have recently been formally differentiated, they have not been empirically distinguished. The aim of this research was to empirically differentiate between parameter learning and structure learning on the basis of how they affect pupil dilation. Participants took part in a within-subject computer-based learning experiment with two phases. In the first phase, participants had to learn the relationship between cues and target stimuli. In the second phase, th...
Notice of republication This article was republished on April 1, 2022, to update the author bylin... more Notice of republication This article was republished on April 1, 2022, to update the author byline and list all members of the group consortium. Please download this article again to view the correct version. The originally published, uncorrected article and the republished, corrected articles are provided here for reference.
The endeavor to understand the human brain has seen more progress in the last few decades than in... more The endeavor to understand the human brain has seen more progress in the last few decades than in the previous two millennia. Still, our understanding of how the human brain relates to behavior in the real world and how this link is modulated by biological, social, and environmental factors is limited. To address this, we designed the Healthy Brain Study (HBS), an interdisciplinary, longitudinal, cohort study based on multidimensional, dynamic assessments in both the laboratory and the real world. Here, we describe the rationale and design of the currently ongoing HBS. The HBS is examining a population-based sample of 1,000 healthy participants (age 30–39) who are thoroughly studied across an entire year. Data are collected through cognitive, affective, behavioral, and physiological testing, neuroimaging, bio-sampling, questionnaires, ecological momentary assessment, and real-world assessments using wearable devices. These data will become an accessible resource for the scientific c...
Within predictive processing learning is construed as Bayesian model updating with the degree of ... more Within predictive processing learning is construed as Bayesian model updating with the degree of certainty for different existing hypotheses changing in light of new evidence. Bayesian model updating, however, cannot explain how new hypotheses are added to a model. Model revision, unlike model updating, makes structural changes to a generative model by altering its causal connections or adding or removing hypotheses. Whilst model updating and model revision have recently been formally differentiated, they have not been empirically distinguished. The aim of this research was to empirically differentiate between model updating and revision on the basis of how they affect prediction errors and predictions over time. To study this, participants took part in a within-subject computer-based learning experiment with two phases: updating and revision. In the updating phase, participants had to predict the relationship between cues and target stimuli and in the revision phase, they had to co...
Behavioral studies have shown that humans account for inertial acceleration in their decisions of... more Behavioral studies have shown that humans account for inertial acceleration in their decisions of hand choice when reaching during body motion. Physiologically, it is unclear at what stage of movement preparation information about body motion is integrated in the process of hand selection. Here, we addressed this question by applying transcranial magnetic stimulation over motor cortex (M1) of human participants who performed a preferential reach task while they were sinusoidally translated on a linear motion platform. If M1 only represents a read-out of the final hand choice, we expect the body motion not to affect the MEP amplitude. If body motion biases the hand selection process prior to target onset, we expect corticospinal excitability to modulate with the phase of the motion, with larger MEP amplitudes for phases that show a bias to using the right hand. Behavioral results replicate our earlier findings of a sinusoidal modulation of hand choice bias with motion phase. MEP ampl...
Contemporary theories of motor control have suggested that multiple motor commands compete for ac... more Contemporary theories of motor control have suggested that multiple motor commands compete for action selection. While most of these competitions are completed prior to movement onset, averaged saccadic eye movements that land at an intermediate location between two visual targets are thought to arise when a movement is initiated prior to the resolution of the competition. In contrast, while averaged reach movements have been reported, there is still debate on whether averaged reach movements are the result of a resolved competition between two potential actions or a strategic behavior that optimally incorporates the current task demands. Here, we use a reach version of the paradigm that has previously shown to elicit saccadic averaging to examine whether similar averaging occurs based on neuromuscular activity of an upper limb muscle. On a single trial basis, we observed a temporal evolution of the two competing motor commands during a free-choice reach task to one of two visual ta...
Recent computational theories and behavioral observations suggest that motor learning is supporte... more Recent computational theories and behavioral observations suggest that motor learning is supported by multiple adaptation processes, operating on different timescales, but direct neural evidence is lacking. We tested this hypothesis by applying transcranial magnetic stimulation over motor cortex in 16 human subjects during a validated reach adaptation task. Motor-evoked potentials (MEPs) and cortical silent periods (CSPs) were recorded from the biceps brachii to assess modulations of corticospinal excitability as indices for corticospinal plasticity. Guided by a two-state adaptation model, we show that the MEP reflects an adaptive process that learns quickly but has poor retention, while the CSP correlates with a process that responds more slowly but retains information well. These results provide a physiological link between models of motor learning and distinct changes in corticospinal excitability. Our findings support the relationship between corticospinal gain modulations and t...
For the brain to decide on a reaching movement, it needs to select which hand to use. A number of... more For the brain to decide on a reaching movement, it needs to select which hand to use. A number of body-centered factors affect this decision, such as the anticipated movement costs of each arm, recent choice success, handedness, and task demands. While the position of each hand relative to the target is also known to be an important spatial factor, it is unclear which reference frames coordinate the spatial aspects in the decisions of hand choice. Here we tested the role of gaze- and head-centered reference frames in a hand selection task. With their head and gaze oriented in different directions, we measured hand choice of 19 right-handed subjects instructed to make unimanual reaching movements to targets at various directions relative to their body. Using an adaptive procedure, we determined the target angle that led to equiprobable right/left hand choices. When gaze remained fixed relative to the body this balanced target angle shifted systematically with head orientation, and wh...
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Papers by Luc Selen