Muscular reflex gains reflect changes of mind in reaching.

Decisions for action are accompanied by a continual processing of sensory information, sometimes resulting in a revision of the initial choice, called a change of mind (CoM). Although the motor system is tuned during the formation of a reach decision, it is unclear whether its preparatory state differs between CoM and non-CoM decisions. To test this, participants (n = 14) viewed a random-dot motion (RDM) stimulus of various coherence levels for a random viewing duration. At the onset of a mechanical perturbation that rapidly stretched the pectoralis muscle, they indicated the perceived motion direction by making a reaching movement to one of two targets. Using electromyography (EMG), we quantified the reflex gains of the pectoralis and posterior deltoid muscles. Results show that reflex gains scaled with both the coherence level and the viewing duration of the stimulus. We fit a drift diffusion model (DDM) to the behavioral choices. The decision variable (DV), derived from the DDM, correlated well with the measured reflex gain at the single-trial level. However, when matched on DV magnitude, reflex gains were significantly lower in CoM than non-CoM trials. We conclude that the internal state of the motor system, as measured by the spinal reflexes, reflects the continual deliberation on sensory evidence for action selection, including the postdecisional evidence that can lead to a change of mind.NEW & NOTEWORTHY Using behavioral findings, EMG, and computational modeling, we show that not only the perceptual decision outcome but also the accumulating evidence for that outcome is continuously sent to the relevant muscles. Moreover, we show that an upcoming change of mind can be detected in the motor periphery, suggesting that a correlate of the internal decision making process is being sent along.

Late integration of vision and proprioception during perturbed reaches.

The motor system corrects rapidly, but selectively, for perturbations to ongoing reaching movements, depending on the constraints of the task. To account for such sophistication, it has been postulated that corrections are based on an estimated limb state that integrates all sensory changes caused by the perturbation, taking into account their processing delays. Here, we asked if information from different sensory modalities is integrated immediately or processed separately in the early phase of a response. We perturbed the estimated state of the limb with both unimodal and bimodal visual and proprioceptive perturbations without changing the actual limb state. For visual perturbations, a cursor representing the hand was shifted to the left or the right relative to the true hand location. For proprioceptive perturbations, the biceps or triceps muscles were vibrated, which induced illusory limb-state changes to the right or the left. In the bimodal condition, the perturbations to vision and proprioception were either congruent or incongruent in their directions. Response latencies show that it takes ∼100 ms longer to respond to unimodal visual perturbations than to unimodal proprioceptive perturbations. Responses to bimodal perturbations show that it takes an additional ∼100 ms beyond the response to unimodal visual perturbations for intermodal consistency to impact the response. These results suggest that visual and proprioceptive signals are initially processed separately for state estimation and only combined at the level of the limb's motor output, instead of being immediately integrated into a single state estimate of the limb.NEW & NOTEWORTHY Both visual and proprioceptive signals provide information about arm state during reaching. By perturbing the perceived, but not the actual, position of the hand in both modalities using visual disturbances and muscle vibration, we examined multimodal integration and state estimation during reaching. Our results suggest that the early reach corrections are based on separate state estimates from the two sensory modalities and only later are based on a combined state estimate.

Single-Trial Dynamics of Competing Reach Plans in the Human Motor Periphery.

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 STATEMENT Contemporary 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.

Plan- versus motion-referenced generalization of fast and slow processes in reach adaptation.

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 plan-referenced 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 11 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.NEW & NOTEWORTHY Error-based reach adaptation can be modeled by fast and slow adaptive processes. Using a spontaneous recovery paradigm and model-based analyses, we tested how these processes generalize during force-field reach adaptation. Depending on whether the fast and slow adaptive processes operate by crediting the planned or actual motion, the model predicts distinct contributions of them to the overall generalization function. We show that human participants fall within a continuum of evidence for plan-referenced to motion-referenced updating.

Abrupt, but not gradual, motor adaptation biases saccadic target selection.

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 toward 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.NEW & NOTEWORTHY Decisions between potential motor actions are influenced by their costs, but costs change when movements are adapted. Using a saccadic target selection task, we show that target preference shifts after abrupt, but not after gradual adaptation. We suggest that this difference emerges because abrupt adaptation results in target remapping, and thus directly influences cost calculations, whereas gradual adaptation is mainly driven by corrections to a forward model that is not involved in cost calculations.

Predictive steering: integration of artificial motor signals in self-motion estimation.

The brain's computations for active and passive self-motion estimation can be unified with a single model that optimally combines vestibular and visual signals with sensory predictions based on efference copies. It is unknown whether this theoretical framework also applies to the integration of artificial motor signals, such as those that occur when driving a car, or whether self-motion estimation in this situation relies on sole feedback control. Here, we examined if training humans to control a self-motion platform leads to the construction of an accurate internal model of the mapping between the steering movement and the vestibular reafference. Participants (n = 15) sat on a linear motion platform and actively controlled the platform's velocity using a steering wheel to translate their body to a memorized visual target (motion condition). We compared their steering behavior to that of participants (n = 15) who remained stationary and instead aligned a nonvisible line with the target (stationary condition). To probe learning, the gain between the steering wheel angle and the platform or line velocity changed abruptly twice during the experiment. These gain changes were virtually undetectable in the displacement error in the motion condition, whereas clear deviations were observed in the stationary condition, showing that participants in the motion condition made within-trial changes to their steering behavior. We conclude that vestibular feedback allows not only the online control of steering but also a rapid adaptation to the gain changes to update the brain's internal model of the mapping between the steering movement and the vestibular reafference.NEW & NOTEWORTHY Perception of self-motion is known to depend on the integration of sensory signals and, when the motion is self-generated, the predicted sensory reafference based on motor efference copies. Here we show, using a closed-loop steering experiment with a direct coupling between the steering movement and the vestibular self-motion feedback, that humans are also able to integrate artificial motor signals, like the motor signals that occur when driving a car.

Exploration of sensory-motor tradeoff behavior in Parkinson's disease.

While slowness of movement is an obligatory characteristic of Parkinson's disease (PD), there are conditions in which patients move uncharacteristically fast, attributed to deficient motor inhibition. Here we investigate deficient inhibition in an optimal sensory-motor integration framework, using a game in which subjects used a paddle to catch a virtual ball. Display of the ball was extinguished as soon as the catching movement started, segregating the task into a sensing and acting phase. We analyzed the behavior of 9 PD patients (ON medication) and 10 age-matched controls (HC). The switching times (between sensing and acting phase) were compared to the predicted optimal switching time, based on the individual estimates of sensory and motor uncertainties. The comparison showed that deviation from predicted optimal switching times were similar between groups. However, PD patients showed a weaker correlation between variability in switching time and sensory-motor uncertainty, indicating a reduced propensity to generate exploratory behavior for optimizing goal-directed movements. Analysis of the movement kinematics revealed that PD patients, compared to controls, used a lower peak velocity of the paddle and intercepted the ball with greater velocity. Adjusting the trial duration to the time for the paddle to stop moving, we found that PD patients spent a smaller proportion of the trial duration for observing the ball. Altogether, the results do not show the premature movement initiation and truncated sensory processing that we predicted to ensue from deficient inhibition in PD.

Assessing corticospinal excitability and reaching hand choice during whole body motion.

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 with the process of hand selection. Here, we addressed this question by applying transcranial magnetic stimulation over left 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 motor-evoked potential (MEP) amplitude. If body motion biases the hand selection process before target onset, we expect corticospinal excitability to be influenced by 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 amplitudes also show a sinusoidal modulation with motion phase, suggesting that body motion influences corticospinal excitability, which may ultimately reflect changes of hand preference. The modulation being present before target onset suggests that competition between hands is represented throughout the corticospinal tract. Its phase relationship with the motion profile indicates that other processes after target onset take up time until the hand selection process has been completely resolved, and the reach is initiated.NEW & NOTEWORTHY 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.

Single versus dual-rate learning when exposed to Coriolis forces during reaching movements.

When we reach for an object during a passive whole body rotation, a tangential Coriolis force is generated on the arm. Yet, within a few trials, the brain adapts to this force so it does not disrupt the reach. Is this adaptation governed by a single-rate or dual-rate learning process? Here, guided by state-space modeling, we studied human reach adaptation in a fully-enclosed rotating room. After 90 pre-rotation reaches (baseline), participants were trained to make 240 to-and-fro reaches while the room rotated at 10 rpm (block A), then performed 6 reaches under opposite room rotation (block B), and subsequently made 100 post-rotation reaches (washout). A control group performed the same paradigm, but without the reaches during rotation block B. Single-rate and dual-rate models can be best dissociated if there would be full un-learning of compensation A during block B, but minimal learning of B. From the perspective of a dual-rate model, the un-learning observed in block B would mainly be caused by the faster state, such that the washout reaches would show retention effects of the slower state, called spontaneous recovery. Alternatively, following a single-rate model, the same state would govern the learning in block A and un-learning in block B, such that the washout reaches mimic the baseline reaches. Our results do not provide clear signs of spontaneous recovery in the washout reaches. Model fits further show that a single-rate process outperformed a dual-rate process. We suggest that a single-rate process underlies Coriolis force reach adaptation, perhaps because these forces relate to familiar body dynamics and are assigned to an internal cause.

Transformation of vestibular signals for the decisions of hand choice during whole body motion.

In daily life, we frequently reach toward objects while our body is in motion. We have recently shown that body accelerations influence the decision of which hand to use for the reach, possibly by modulating the body-centered computations of the expected reach costs. However, head orientation relative to the body was not manipulated, and hence it remains unclear whether vestibular signals contribute in their head-based sensory frame or in a transformed body-centered reference frame to these cost calculations. To test this, subjects performed a preferential reaching task to targets at various directions while they were sinusoidally translated along the lateral body axis, with their head either aligned with the body (straight ahead) or rotated 18° to the left. As a measure of hand preference, we determined the target direction that resulted in equiprobable right/left-hand choices. Results show that head orientation affects this balanced target angle when the body is stationary but does not further modulate hand preference when the body is in motion. Furthermore, reaction and movement times were larger for reaches to the balanced target angle, resembling a competitive selection process, and were modulated by head orientation when the body was stationary. During body translation, reaction and movement times depended on the phase of the motion, but this phase-dependent modulation had no interaction with head orientation. We conclude that the brain transforms vestibular signals to body-centered coordinates at the early stage of reach planning, when the decision of hand choice is computed. NEW & NOTEWORTHY The brain takes inertial acceleration into account in computing the anticipated biomechanical costs that guide hand selection during whole body motion. Whereas these costs are defined in a body-centered, muscle-based reference frame, the otoliths detect the inertial acceleration in head-centered coordinates. By systematically manipulating head position relative to the body, we show that the brain transforms otolith signals into body-centered coordinates at an early stage of reach planning, i.e., before the decision of hand choice is computed.

Visual orientation uncertainty in the rod-and-frame illusion.

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 (-17.5°, 0°, 17.5°). 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.

Age-related reweighting of visual and vestibular cues for vertical perception.

As we age, the acuity of our sensory organs declines, which may affect our lifestyle. Sensory deterioration in the vestibular system is typically bilateral and gradual, and could lead to problems with balance and spatial orientation. To compensate for the sensory deterioration, it has been suggested that the brain reweights the sensory information sources according to their relative noise characteristics. For rehabilitation and training programs, it is important to understand the consequences of this reweighting, preferably at the individual subject level. We psychometrically examined the age-dependent reweighting of visual and vestibular cues used in spatial orientation in a group of 32 subjects (age range: 19-76 yr). We asked subjects to indicate the orientation of a line (clockwise or counterclockwise relative to the gravitational vertical) presented within an oriented square visual frame when seated upright or with their head tilted 30° relative to the body. Results show that subjects' vertical perception is biased by the orientation of the visual frame. Both the magnitude of this bias and response variability become larger with increasing age. Deducing the underlying sensory noise characteristics, using Bayesian inference, suggests an age-dependent reweighting of sensory information, with an increasing weight of the visual contextual information. Further scrutiny of the model suggests that this shift in sensory weights is the result of an increase in the noise of the vestibular signal. Our approach quantifies how noise properties of visual and vestibular systems change over the life span, which helps to understand the aging process at the neurocomputational level. NEW & NOTEWORTHY Perception of visual vertical involves a weighted fusion of visual and vestibular tilt cues. Using a Bayesian approach and experimental psychophysics, we quantify how this fusion process changes with age. We show that, with age, the vestibular information is down-weighted whereas the visual weight is increased. This shift in sensory reweighting is primarily due to an age-related increase of the noise of vestibular signals.

Task-dependent responses to muscle vibration during reaching.

Feedback corrections in reaching have been shown to be task-dependent for proprioceptive, visual and vestibular perturbations, in line with predictions from optimal feedback control theory. Mechanical perturbations have been used to elicit proprioceptive errors, but have the drawback to actively alter the limb's trajectory, making it nontrivial to dissociate the subject's compensatory response from the perturbation itself. In contrast, muscle vibration provides an alternative tool to perturb the muscle afferents without changing the hands trajectory, inducing only changes in the estimated, but not the actual, limb position and velocity. Here, we investigate whether upper-arm muscle vibration is sufficient to evoke task-dependent feedback corrections during goal-directed reaching to a narrow versus a wide target. Our main result is that for vibration of biceps and triceps, compensatory responses were down-regulated for the wide compared to the narrow target. The earliest detectable difference between these target-specific corrections is at about 100 ms, likely reflecting a task-dependent feedback control policy rather than a voluntary response.

Bayesian adaptive stimulus selection for dissociating models of psychophysical data.

Comparing models facilitates testing different hypotheses regarding the computational basis of perception and action. Effective model comparison requires stimuli for which models make different predictions. Typically, experiments use a predetermined set of stimuli or sample stimuli randomly. Both methods have limitations; a predetermined set may not contain stimuli that dissociate the models, whereas random sampling may be inefficient. To overcome these limitations, we expanded the psi-algorithm (Kontsevich & Tyler, 1999) from estimating the parameters of a psychometric curve to distinguishing models. To test our algorithm, we applied it to two distinct problems. First, we investigated dissociating sensory noise models. We simulated ideal observers with different noise models performing a two-alternative forced-choice task. Stimuli were selected randomly or using our algorithm. We found using our algorithm improved the accuracy of model comparison. We also validated the algorithm in subjects by inferring which noise model underlies speed perception. Our algorithm converged quickly to the model previously proposed (Stocker & Simoncelli, 2006), whereas if stimuli were selected randomly, model probabilities separated slower and sometimes supported alternative models. Second, we applied our algorithm to a different problem-comparing models of target selection under body acceleration. Previous work found target choice preference is modulated by whole body acceleration (Rincon-Gonzalez et al., 2016). However, the effect is subtle, making model comparison difficult. We show that selecting stimuli adaptively could have led to stronger conclusions in model comparison. We conclude that our technique is more efficient and more reliable than current methods of stimulus selection for dissociating models.

Corticospinal correlates of fast and slow adaptive processes in motor learning.

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 the adaptive processes in motor learning. NEW & NOTEWORTHY Computational theories and behavioral observations suggest that motor learning is supported by multiple adaptation processes, but direct neural evidence is lacking. We tested this hypothesis by applying transcranial magnetic stimulation over human motor cortex during a reach adaptation task. Guided by a two-state adaptation model, we show that the motor-evoked potential reflects a process that adapts and decays quickly, whereas the cortical silent period reflects slow adaptation and decay.

Uncertainty modulated exploration in the trade-off between sensing and acting.

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.

Psychophysical Evaluation of Sensory Reweighting in Bilateral Vestibulopathy.

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.

Reference frames in the decisions of hand choice.

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 when head orientation remained fixed this choice measure shifted with gaze. These results suggest that a mixture of head- and gaze-centered reference frames is involved in the spatially guided decisions of hand choice, perhaps to flexibly bind this process to the mechanisms of target selection. NEW & NOTEWORTHY Decisions of target and hand choice are fundamental aspects of human reaching movements. While the reference frames involved in target choice have been identified, it is unclear which reference frames are involved in hand selection. We tested the role of gaze- and head-centered reference frames in a hand selection task. Findings emphasize the role of both spatial reference frames in the decisions of hand choice, in addition to known body-centered computations such anticipated movement costs and handedness.

Bayesian quantification of sensory reweighting in a familial bilateral vestibular disorder (DFNA9).

DFNA9 is a rare progressive autosomal dominantly inherited vestibulo-cochlear disorder, resulting in a homogeneous group of patients with hearing impairment and bilateral vestibular function loss. These patients suffer from a deteriorated sense of spatial orientation, leading to balance problems in darkness, especially on irregular surfaces. Both behavioral and functional imaging studies suggest that the remaining sensory cues could compensate for the loss of vestibular information. A thorough model-based quantification of this reweighting in individual patients is, however, missing. Here we psychometrically examined the individual patient's sensory reweighting of these cues after complete vestibular loss. We asked a group of DFNA9 patients and healthy control subjects to judge the orientation (clockwise or counterclockwise relative to gravity) of a rod presented within an oriented square frame (rod-in-frame task) in three different head-on-body tilt conditions. Our results show a cyclical frame-induced bias in perceived gravity direction across a 90° range of frame orientations. The magnitude of this bias was significantly increased in the patients compared with the healthy control subjects. Response variability, which increased with head-on-body tilt, was also larger for the patients. Reverse engineering of the underlying signal properties, using Bayesian inference principles, suggests a reweighting of sensory signals, with an increase in visual weight of 20-40% in the patients. Our approach of combining psychophysics and Bayesian reverse engineering is the first to quantify the weights associated with the different sensory modalities at an individual patient level, which could make it possible to develop personal rehabilitation programs based on the patient's sensory weight distribution. NEW & NOTEWORTHY It has been suggested that patients with vestibular deficits can compensate for this loss by increasing reliance on other sensory cues, although an actual quantification of this reweighting is lacking. We combine experimental psychophysics with a reverse engineering approach based on Bayesian inference principles to quantify sensory reweighting in individual vestibular patients. We discuss the suitability of this approach for developing personal rehabilitation programs based on the patient's sensory weight distribution.

Flexible Visuomotor Associations in Touchscreen Control.

To move real objects, our hand needs to get in direct physical contact with the object. However, this is not necessarily the case when interacting with virtual objects, for example when displacing objects on tablets by swipe movements. Here, we performed two experiments to study the behavioral strategies of these movements, examining how visual information about the virtual object is mapped into a swipe that moves the object into a goal location. In the first experiment, we investigated how swiping behavior depends on whether objects were located within or outside the swiping workspace. Results show that participants do not start the swipe movement by placing their finger on the virtual object, as they do when reaching to real objects, but rather keep a systematic distance between the object location and the initial swipe location. This mismatch, which was experimentally imposed by placing the object outside the workspace, also occurred when the object was within the workspace. In the second experiment, we investigated which factors determine this mismatch by systematically manipulating the initial hand location, the location of the object and the location of the goal. Dimensionality reduction of the data showed that three factors are taken into account when participants choose the initial swipe location: the expected total movement distance, the distance between their finger on the screen and the object, and a preference not to cover the object. The weight given to each factor differed among individuals. These results delineate, for the first time, the flexibility of visuomotor associations in the virtual world.