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Medendorp, W. P.

Publications and source records attributed to Medendorp, W. P..

29 records · Page 2Linked to original sources

Natural statistics of head roll: implications for Bayesian inference in spatial orientation

We previously proposed a Bayesian model of multisensory integration in spatial orientation (1). Using a Gaussian prior, centered on an upright head orientation, this model could explain various perceptual observations in roll-tilted participants, such as the subjective visual vertical, the subjective body tilt (1), the rod-and-frame effect (2), as well as their clinical (3) and age-related deficits (4). Because it is generally assumed that the prior reflects an accumulated history of previous head orientations, and recent work on natural head motion suggests non-Gaussian statistics, we examined how the model would perform with a non-Gaussian prior. In the present study, we first experimentally generalized the previous observations in showing that also the natural statistics of head orientation are characterized by long tails, best quantified as a t-location-scale distribution. Next, we compared the performance of the Bayesian model and various model variants using such a t-distributed prior to the original model with the Gaussian prior on their accounts of previously published data of the subjective visual vertical and subjective body tilt tasks. All of these variants performed substantially worse than the original model, suggesting a special value of the Gaussian prior. We provide computational and neurophysiological reasons for the implementation of such a prior, in terms of its associated precision-accuracy trade-off in vertical perception across the tilt range. New & NoteworthyIt has been argued that the brain uses Bayesian computations to process multiple sensory cues in vertical perception, including a prior centered on upright head orientation which is usually taken to be Gaussian. Here, we show that non-Gaussian prior distributions, although more akin to the statistics of head orientation during natural activities, provide a much worse explanation of such perceptual observations than a Gaussian prior.

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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 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.

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

The brains 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 motor efference copies. It is unknown whether this theoretical framework also applies to the integration of artificial motor signals, like the motor signals that occur when driving a car. Here, we examined if training humans to control a self-motion platform would lead to the construction of an accurate internal model of the mapping between the steering movement and the vestibular reafference. Participants (n = 15) were seated on a linear motion platform and actively controlled the platforms velocity using a steering wheel to translate their body to a memorized visual target location (Motion condition). We compared their steering behavior to that of participants (n = 15) who remained stationary and instead aligned a non-visible line with the target (Stationary condition). To probe learning, the gain between the steering wheel angle and the platform velocity 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. These results show that participants in the Motion condition made within-trial changes to their steering behavior immediately after the gain changes. This suggests that they continuously compared the vestibular reafference to internal predictions, and thus employed and updated an internal forward model of the mapping between the steering movement and the vestibular reafference. New & NoteworthyPerception 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.

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A horizon for haptic perception

The spatial limits of sensory acquisition (its sensory horizon) is a fundamental property of any sensorimotor system. In the present study, we sought to determine the sensory horizon for the human haptic modality. At first blush, it seems obvious that the haptic system is bounded by the space where the body can interact with the environment (e.g., the arm span). However, the human somatosensory system is exquisitely tuned to sensing with tools--blind-cane navigation being a classic example of this. The horizon of haptic perception therefore extends beyond body space, but to what extent is unknown. We first used neuromechanical modelling to determine the theoretical horizon, which we pinpointed as six meters. We then used a psychophysical localization paradigm to behaviorally confirm that humans can haptically localize objects using a six-meter rod. This finding underscores the incredibly flexibility of the brains sensorimotor representations, as they can be adapted to sense with an object many times longer than the users own body.

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Even well-practiced movements benefit from repetition

Professional golf players spend years practicing, but will still perform one or two practice swings without a ball before executing the actual swing. Why do they do this? In this study we tested the hypothesis that repeating a well-practiced movement leads to a reduction of movement variability. To operationalize this hypothesis, participants were tested in a center- out reaching task with four different targets, on four different days. To probe the effect of repetition they performed random sequences from one to six movements to the same target. Our findings show that, with repetition, movements are not only initiated earlier but their variability is reduced across the entire movement trajectory. Furthermore, this effect is present within and across the four sessions. Together, our results suggest that movement repetition changes the tradeoff between movement initiation and movement precision. New & NoteworthyProfessional athletes practice movements that they have performed thousands of times in training just before it is their turn in a game. Why do they do this? Our results indicate that both initial and endpoint variability reduce with repetition in a short sequence of reaching movements. This means that even well-practiced movements benefit from practice.

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A somatosensory computation that unifies bodies and tools

It is often claimed that tools are embodied by the user, but whether the brain actually repurposes its body-based computations to perform similar tasks with tools is not known. A fundamental computation for localizing touch on the body is trilateration. Here, the location of touch on a limb is computed by integrating estimates of the distance between sensory input and its boundaries (e.g., elbow and wrist of the forearm). As evidence of this computational mechanism, tactile localization on a limb is most precise near its boundaries and lowest in the middle. We show that the brain repurposes trilateration to localize touch on a tool. In a large sample of participants, we found that localizing touch on a tool produced the signature of trilateration, with highest precision close to the base and tip of the tool. A computational model of trilateration provided a good fit to the observed localization behavior. To further demonstrate the computational plausibility of repurposing trilateration, we implemented it in a three-layer neural network that was based on principles of probabilistic population coding. This network determined hit location in tool-centered coordinates by using a tools unique pattern of vibrations when contacting an object. Simulations demonstrated the expected signature of trilateration, in line with the behavioral patterns. Our results have important implications for how trilateration may be implemented by somatosensory neural populations. We conclude that trilateration is a fundamental spatial computation that unifies limbs and tools.

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Beta-band desynchronization reflects uncertainty in effector selection during motor planning

While beta-band activity during motor planning is known to be modulated by uncertainty about where to act, less is known about its modulations to uncertainty about how to act. To investigate this issue, we recorded oscillatory brain activity with EEG while human participants (n = 17) performed a hand choice reaching task. The reaching hand was either predetermined or of participants choice, and the target was close to one of the two hands or at about equal distance from both. To measure neural activity in a motion-artifact-free time window, the location of the upcoming target was cued 1000-1500 ms before the presentation of the target, whereby the cue was valid in 50% of trials. As evidence for motor planning during the cueing phase, behavioral observations showed that the cue affected later hand choice. Furthermore, reaction times were longer in the choice than in the predetermined trials, supporting the notion of a competitive process for hand selection. Modulations of beta-band power over central cortical regions, but not alpha-band or theta-band power, were in line with these observations. During the cueing period, reaches in predetermined trials were preceded by larger decreases in beta-band power than reaches in choice trials. Cue direction did not affect reaction times or beta-band power, which may be due to the cue being invalid in 50% of trials, retaining effector uncertainty during motor planning. Our findings suggest that effector uncertainty, similar to target uncertainty, selectively modulates beta-band power during motor planning. New & NoteworthyWhile reach-related beta-band power in central cortical areas is known to modulate with the number of potential targets, here we show, using a cueing paradigm, that the power in this frequency band, but not in the alpha or theta-band, is also modulated by the uncertainty of which hand to use. This finding supports the notion that multiple possible effector-specific actions can be specified in parallel up to the level of motor preparation.

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Corticospinal correlates of hand preference for reaching 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 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 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 prior to target onset suggests that competition between hands is represented throughout the corticospinal tract. Its phase relationship with the motion profile suggests that other processes after target onset take up time until the hand selection process has been completely resolved, and the reach is initiated. We conclude that the corticospinal correlates of hand preference are modulated by body motion.

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A neural surveyor in somatosensory cortex

Perhaps the most recognizable sensory map in all of neuroscience is the somatosensory homunculus. Though it seems straightforward, this simple representation belies the complex link between an activation in a somatotopic map and the associated touch location on the body. Any isolated activation is spatially ambiguous without a neural decoder that can read its position within the entire map, but how this is computed by neural networks is unknown. We propose that the somatosensory system implements multilateration, a common computation used by surveying and GPS systems to localize objects. Specifically, to decode touch location on the body, multilateration estimates the relative distance between the afferent input and the boundaries of a body part (e.g., the joints of a limb). We show that a simple feedforward neural network, which captures several fundamental receptive field properties of cortical somatosensory neurons, can implement a Bayes-optimal multilateral computation. Simulations demonstrated that this decoder produced a pattern of localization variability between two boundaries that was unique to multilateration. Finally, we identify this computational signature of multilateration in actual psychophysical experiments, suggesting that it is a candidate computational mechanism underlying tactile localization.

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External location of touch is constructed post-hoc based on limb choice

When humans indicate on which hand a tactile stimulus occurred, they often err when their hands are crossed. This finding seemingly supports the view that the automatically determined touch location in external space affects limb assignment: the crossed right hand is localized in left space, and this conflict presumably provokes hand assignment errors. Here, participants judged on which hand the first of two stimuli, presented during a bimanual movement, had occurred, and then indicated its external location by a reach-to-point movement. When participants incorrectly chose the hand stimulated second, they pointed to where that hand had been at the correct, first time point, though no stimulus had occurred at that location. This behavior suggests that stimulus localization depended on hand assignment, not vice versa. It is, thus, incompatible with the notion of automatic computation of external stimulus location upon occurrence. Instead, humans construct external touch location post-hoc and on demand.

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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 2-afc 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.\n\nData AvailabilityAll data and code will be posted on our institutional repository system following acceptance. In the meantime feel free to contact the authors if you would like any of the code.

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