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Biomedical subjects

M Kawato

Publications and source records attributed to M Kawato.

At least 37 records · Page 2Linked to original sources

Using motor tasks to quantitatively judge 3-D surface curvatures.

The primary objective of this study was to quantitatively investigate the human perception of surface curvature by using virtual surfaces and motor tasks along with data analysis methods to estimate surface curvature from drawing movements. Three psychophysical experiments were conducted. In Experiment 1, we looked at subjects' sensitivity to the curvature of a curve lying on a surface and changes in the curvature as defined by Euler's formula, which relates maximum and minimum principal curvatures and their directions. Regardless of direction and surface shape (elliptic and hyperbolic), subjects could report the curvature of a curve lying on a surface through a drawing task. In addition, multiple curves drawn by subjects were used to reconstruct the surface. These reconstructed surfaces could be better accounted for by analysis that treated the drawing data as a set of curvatures rather than as a set of depths. A pointing task was utilized in Experiment 2, and subjects could report principal curvature directions of a surface rather precisely and consistently when the difference between principal curvatures was sufficiently large, but performance was poor for the direction of zero curvature (asymptotic direction) on a hyperbolic surface. In Experiment 3, it was discovered that sensitivity to the sign of curvature was different for perceptual judgments and motor responses, and there was also a difference for that of a curve itself and the same curve embedded in a surface. These findings suggest that humans are sensitive to relative changes in curvature and are able to comprehend quantitative surface curvature for some motor tasks.

Adult↗

Internal models for motor control.

The process of moving the hand to a target in space involves a series of sensorimotor transformations that translate visual and other sensory information about the location of the target object and the limbs into a set of motor commands that will bring the hand to the desired position. Recent work at various laboratories has provided strong support for the hypothesis that the CNS learns and maintains internal models of sensorimotor transformations. An internal model is a neural system that mimics the behaviour of the sensorimotor system and objects in the external environment. Internal models enable the CNS to predict the consequences of motor commands and to determine the motor commands required to perform specific tasks. In this chapter, we first summarize recent computational, behavioural and neurophysiological studies that address the theoretical necessity of internal models, the locations of internal models, and the neural mechanism for acquiring internal models through learning. Then, we propose a new computational model of multiple internal models.

Animals↗

Multiple paired forward and inverse models for motor control.

Humans demonstrate a remarkable ability to generate accurate and appropriate motor behavior under many different and often uncertain environmental conditions. In this paper, we propose a modular approach to such motor learning and control. We review the behavioral evidence and benefits of modularity, and propose a new architecture based on multiple pairs of inverse (controller) and forward (predictor) models. Within each pair, the inverse and forward models are tightly coupled both during their acquisition, through motor learning, and use, during which the forward models determine the contribution of each inverse model's output to the final motor command. This architecture can simultaneously learn the multiple inverse models necessary for control as well as how to select the inverse models appropriate for a given environment. Finally, we describe specific predictions of the model, which can be tested experimentally.

Journal Article↗

Adaptive internal model of intrinsic kinematics involved in learning an aiming task.

The elbow-joint angle and the shoulder-joint angle of participants aiming at targets were multiplied in an experiment that used a position-recording system and a cathode-ray tube screen. The linear transformation in joint angles (intrinsic coordinates) corresponded to a nonlinear transformation between the hand coordinates and the screen coordinates (extrinsic coordinates). We examined whether participants could learn this transformation in the intrinsic coordinates or in the extrinsic coordinates by investigating intermanual (between-hands) transfer under an intrinsically consistent condition and an extrinsically consistent condition. Positive intermanual transfer was observed in the former condition but not in the latter condition. Results suggest that participants can learn the linear transformation in joint angles under the intrinsic coordinates and that the central nervous system adaptively represents the intrinsic kinematics.

Adaptation, Psychological↗

Role of the cerebellum in reaching movements in humans. I. Distributed inverse dynamics control.

This study focuses on the role of the motor cortex, the spinal cord and the cerebellum in the dynamics stage of the control of arm movement. Currently, two classes of models have been proposed for the neural control of movements, namely the virtual trajectory control hypothesis and the acquisition of internal models of the motor apparatus hypothesis. In the present study, we expand the virtual trajectory model to whole arm reaching movements. This expanded model accurately reproduced slow movements, but faster reaching movements deviated significantly from the planned trajectories, indicating that for fast movements, this model was not sufficient. These results led us to propose a new distributed functional model consistent with behavioural, anatomical and neurophysiological data, which takes into account arm muscles, spinal cord, motor cortex and cerebellum and is consistent with the view that the central nervous system acquires a distributed inverse dynamics model of the arm. Previous studies indicated that the cerebellum compensates for the interaction forces that arise during reaching movements. We show here how the cerebellum may increase the accuracy of reaching movements by compensating for the interaction torques by learning a portion of an inverse dynamics model that refines a basic inverse model in the motor cortex and spinal cord.

Arm↗

Role of the cerebellum in reaching movements in humans. II. A neural model of the intermediate cerebellum.

The cerebellum is essential for the control of multijoint movements; when the cerebellum is lesioned, the performance error is more than the summed errors produced by single joints. In the companion paper (Schweighofer et al., 1998), a functional anatomical model for visually guided arm movement was proposed. The model comprised a basic feedforward/feedback controller with realistic transmission delays and was connected to a two-link, six-muscle, planar arm. In the present study, we examined the role of the cerebellum in reaching movements by embedding a novel, detailed cerebellar neural network in this functional control model. We could derive realistic cerebellar inputs and the role of the cerebellum in learning to control the arm was assessed. This cerebellar network learned the part of the inverse dynamics of the arm not provided by the basic feedforward/feedback controller. Despite realistically low inferior olive firing rates and noisy mossy fibre inputs, the model could reduce the error between intended and planned movements. The responses of the different cell groups were comparable to those of biological cell groups. In particular, the modelled Purkinje cells exhibited directional tuning after learning and the parallel fibres, due to their length, provide Purkinje cells with the input required for this coordination task. The inferior olive responses contained two different components; the earlier response, locked to movement onset, was always present and the later response disappeared after learning. These results support the theory that the cerebellum is involved in motor learning.

Arm↗

Temporal firing patterns of Purkinje cells in the cerebellar ventral paraflocculus during ocular following responses in monkeys I. Simple spikes.

The simple-spike firing frequency of 30 Purkinje cells (P cells) in the ventral paraflocculus (VPFL) of alert monkeys was studied in relation to vertical slow eye movements, termed ocular following response (OFR), induced by large-field visual motions of different velocities and durations. To quantitatively analyze the relationship between eye movement and firing frequency, an inverse dynamics representation of the eye movement was used for reconstructing the temporal waveform of firing. Coefficients of eye-acceleration, velocity, and position, bias, and time lag between firing and eye movement were estimated by least-square error method. In the regression analyses for each stimulus condition, 86% (146/170) of the well-modulated temporal firing patterns taken from those 30 P cells were reconstructed successfully from eye movement. The model with acceleration, velocity, and position terms, which we used, was shown as the best among several potential models by Cp statistics, consistent with t-test of significance of each term. Reliable coefficients were obtained from 75% (109/146) of the well-reconstructed firing patterns of 28 cells among 30. The estimated coefficients were larger (statistically significant) for slow stimuli than for fast stimuli, suggesting changes in sensitivities under different conditions. However, firing patterns of each cell under several different conditions were frequently well reconstructed by an inverse dynamics representation with a single set of coefficients (13 cells among 21). This indicates that the relationships between P cell firing and OFR are roughly linear in those stimulus ranges. The estimated coefficients for acceleration and velocity suggested that the VPFL P cells properly encode the dynamic components of the motor command during vertical OFR. As for the positional component, however, these P cells are correlated with eye movement in the opposite direction. In the regression analysis without positional component, remarkable differences between observed and reconstructed firing patterns were noted especially in the initial phase of the movements, indicating that the negative positional component was not negligible during OFR. Thus we conclude that, during OFR, the VPFL P cells cannot provide the necessary final motor command, and other brain regions, downstream neural structures, or other types of P cells must provide lacking position-dependent motor commands. This finding about the negative correlation with the position is in the opposite sign with previous studies obtained from the fixation and the smooth pursuit movement. From these comparisons, how the VPFL contributes to a part of the final motor command or how other brain regions complement the VPFL is suggested to be different for early and late phases of the movements.

Action Potentials↗

Temporal firing patterns of Purkinje cells in the cerebellar ventral paraflocculus during ocular following responses in monkeys II. Complex spikes.

Many theories of cerebellar motor learning propose that complex spikes (CS) provide essential error signals for learning and modulate parallel fiber inputs that generate simple spikes (SS). These theories, however, do not satisfactorily specify what modality is represented by CS or how information is conveyed by the ultra-low CS firing rate (1 Hz). To further examine the function of CS and the relationship between CS and SS in the cerebellum, CS and SS were recorded in the ventral paraflocculus (VPFL) of awake monkeys during ocular following responses (OFR). In addition, a new statistical method using a generalized linear model of firing probability based on a binomial distribution of the spike count was developed for analysis of the ultra-low CS firing rate. The results of the present study showed that the spatial coordinates of CS were aligned with those of SS and the speed-tuning properties of CS and SS were more linear for eye movement than retinal slip velocity, indicating that CS contain a motor component in addition to the sensory component identified in previous studies. The generalized linear model to reproduce firing probability confirmed these results, demonstrating that CS conveyed high-frequency information with its ultra-low firing frequency and conveyed both sensory and motor information. Although the temporal patterns of the CS were similar to those of the SS when the sign was reversed and magnitude was amplified approximately 50 times, the velocity/acceleration coefficient ratio of the eye movement model, an aspect of the CS temporal firing profile, was less than that of the SS, suggesting that CS were more sensory in nature than SS. A cross-correlation analysis of SS that are triggered by CS revealed that short-term modulation, that is, the brief pause in SS caused by CS, does not account for the reciprocal modulation of SS and CS. The results also showed that three major aspects of the CS and SS individual cell firing characteristics were negatively correlated on a cell-to-cell basis: the preferred direction of stimulus motion, the mean percent change in firing rate induced by upward stimulus motion, and patterns of temporal firing probability. These results suggest that CS may contribute to long-term interactions between parallel and climbing fiber inputs, such as long-term depression and/or potentiation.

Action Potentials↗

Ambulatory physical activity monitoring system.

In this study, we have developed an ambulatory behaviour map and physical activity monitoring system by equipping our portable digital biosignal memory device developed previously with a GPS sensor and piezoresistive accelerometers. By this system, we can get the subjects behaviour map, his physical activities and posture changes in daily life.

Behavior↗

Human arm stiffness and equilibrium-point trajectory during multi-joint movement.

By using a newly designed high-performance manipulandum and a new estimation algorithm, we measured human multi-joint arm stiffness parameters during multi-joint point-to-point movements on a horizontal plane. This manipulandum allows us to apply a sufficient perturbation to subject's arm within a brief period during movement. Arm stiffness parameters were reliably estimated using a new algorithm, in which all unknown structural parameters could be estimated independent of arm posture (i.e., constant values under any arm posture). Arm stiffness during transverse movement was considerably greater than that during corresponding posture, but not during a longitudinal movement. Although the ratios of elbow, shoulder, and double-joint stiffness were varied in time, the orientation of stiffness ellipses during the movement did not change much. Equilibrium-point trajectories that were predicted from measured stiffness parameters and actual trajectories were slightly sinusoidally curved in Cartesian space and their velocity profiles were quite different from the velocity profiles of actual hand trajectories. This result contradicts the hypothesis that the brain does not take the dynamics into account in movement control depending on the neuromuscular servo mechanism; rather, it implies that the brain needs to acquire some internal models of controlled objects.

Adult↗

A mathematical model that reproduces vertical ocular following responses from visual stimuli by reproducing the simple spike firing frequency of Purkinje cells in the cerebellum.

A mathematical model that accurately reproduces eye movements from visual stimuli and incorporates intermediate neural signals is useful for quantitative analysis of the neural mechanisms involved in transforming visual stimuli to eye movements. Here we describe a mathematical model consisting of two systems: a non-linear system that relates retinal slip to simple spike firing frequency of Purkinje cells in the ventral paraflocculus (VPFL) and a linear system that relates VPFL simple spike firing frequency to eye movement. This model accurately reproduced the firing frequency of Purkinje cells and ocular following responses from visual stimulation paradigms used in physiological experiments.

Action Potentials↗

Possible explanations for trajectory curvature in multijoint arm movements.

Although the straightness of hand paths is a widely accepted feature of human multijoint reaching movement, detailed examinations have revealed slight curvatures in some regions of the workspace. This observation raises the question of whether planned trajectories are straight or curved. If they are straight, 3 possible factors can explain the observed curvatures: (a) imperfect control, (b) visual distortion, or (c) interaction between straight virtual trajectories and the dynamics of the arm. Participants instructed to generate straight movement paths produced movements much straighter than those generated spontaneously. Participants generated spontaneously curved trajectories in the frontoparallel plane, where visual distortion is not expected. Electromyograms suggested that participants generated straighter paths without an increase in arm stiffness. These findings argue against the 3 factors. It follows that planned trajectories are likely to be curved.

Adult↗

Correspondence in pictorial space.

We have investigated psychophysically determined image correspondences between pairs of photographs of a single three-dimensional (3-D) object in various poses. These correspondences were obtained by presenting the pictures simultaneously, side by side, and letting the subject match a marker in one picture with a marker (under manual control) in the other picture. Between poses, the object was rotated about a fixed vertical axis; thus, the shifts of the veridical correspondences (with respect to the surface of the object) were very nearly horizontal. In fact, the subjects produced appreciable scatter in both horizontal and vertical directions. The scatter in repeated sessions and between data depends on the local (landmarks) and global (interpolation) structure of the pictures. Since the object was fairly smooth (white semigloss finish) and nontextured, the only way to establish the correspondence is by way of the "pictorial relief." The relief is some largely unknown function of the image structure and the observer. Apparently, more immediate entities (e.g., the shading or the contour) cannot be used as such, since they vary with the pose. We compare these data with results obtained with a surface attitude probe on a single picture. We studied various measures of consistency both within a single method and between methods. We found that subjects were confident in establishing correspondences, but results scattered appreciably in a way that depended on both global and local image structure. Correspondence results for various pose angles were mutually very consistent, but only to a minor extent with results of attitude measurements. The main finding was that subjects could establish correspondence on the basis of their 3-D interpretation (pictorial relief), even if the 2-D graytone distributions are quite different.

Form Perception↗

Perception of local orientation from shaded images.

The perception of local orientation from shaded images was examined. In Experiment 1, subjects viewed a boundaryless Gaussian hill and judged local orientation using both a gauge figure and a pointing method. One subject reported an internally consistent surface which was incompatible with the judged light-source direction and model used to generate the image. The remaining subjects reported a surface similar to the generating one, and analysis of their results indicated a contour of zero difference between response and generating slants. This contour of zero slant difference was explored in three subsequent experiments using the pointing technique. These experiments investigated possible influences of luminance artifact (Experiment 2), perception of global orientation (Experiment 3), and self-occluding contours (Experiment 4). All three of these experiments yielded results similar to those of Experiment 1, with distinct contours of zero slant difference. This contour was explored for relationships with the simulated slant of the generating surface and the differential structure of image intensity. This analysis indicated that the contour of zero slant difference was approximately a line of constant slant which shared large regions of adjacency to the zero crossings of the second directional derivative of image intensity.

Adult↗

A theory for cursive handwriting based on the minimization principle.

We propose a trajectory planning and control theory which provides explanations at the computation, algorithm, representation, and hardware levels for continuous movement such as connected cursive handwriting. The hardware is based on our previously proposed forward-inverse-relaxation neural network. Computationally, the optimization principle is the minimum torque-change criterion. At the representation level, hard constraints satisfied by a trajectory are represented as a set of via-points extracted from handwritten characters. Accordingly, we propose a via-point estimation algorithm that estimates via-points by repeating trajectory formation of a character and via-point extraction from the character. It is shown experimentally that for movements with a single via-point target, the via-point estimation algorithm can assign a point near the actual via-point target. Good quantitative agreement is found between human movement data and the trajectories generated by the proposed model.

Algorithms↗

A computational theory for movement pattern recognition based on optimal movement pattern generation.

We have previously proposed an optimal trajectory and control theory for continuous movements, such as reaching or cursive handwriting. According to Marr's three-level description of brain function, our theory can be summarized as follows: (1) The computational theory is the minimum torque-change model; (2) the intermediate representation of a pattern is given as a set of via-points extracted from an example pattern; and (3) algorithm and hardware are provided by FIRM, a neural network that can generate and control minimum torque-change trajectories. In this paper, we propose a computational theory for movement pattern recognition that is based on our theory for optimal movement pattern generation. The three levels of the description of brain function in the recognition theory are tightly coupled with those for pattern generation. In recognition, the generation process and the recognition process are actually two flows of information in opposite directions within a single functional unit. In our theory, if the input movement trajectory data are identical to the optimal movement pattern reconstructed from an intermediate representation of some symbol, the input data are recognized as that symbol. If an error exists between the movement trajectory data and the generated trajectory, the putative symbol is corrected, and the generation is repeated. In particular, we present concrete computational procedures for the recognition of connected cursive handwritten characters, as well as for the estimation of phonemic timing in natural speech. Our most important contribution is to demonstrate the computational realizability for the 'motor theory of movement pattern perception': the movement-pattern recognition process can be realized by actively recruiting the movement-pattern formation process. The way in which the formation process is utilized in pattern recognition in our theory suggests a duality between movement pattern formation and movement pattern perception.

Algorithms↗