Christophe Bourdin

Galvanic vestibular stimulation in humans produces online arm movement deviations when reaching towards memorized visual targets.

Using galvanic vestibular stimulation (GVS), we tested whether a change in vestibular input at the onset of goal-directed arm movements induces deviations in arm trajectory. Eight head-fixed standing subjects were instructed to reach for memorized visual targets in complete darkness. In half of the trials, randomly-selected, a 3 mA bipolar binaural galvanic stimulation of randomly alternating polarity was triggered by the movement onset. Results revealed significant GVS-induced directional shifts of reaching movements towards the anode side. The earliest significant deviations of hand path occurred 240 ms after stimulation onset. The likely goal of these online deviations of arm trajectory was to compensate for a vestibular-evoked apparent change in the spatial relationship between the target and the hand.

Role of sensory information in updating internal models of the effector during arm tracking

This chapter is divided into three main parts. Firstly, on the basis of the literature, we will shortly discuss how the recent introduction of the concept of internal models by Daniel Wolpert and Mitsuo Kawato contributes to a better understanding of what is motor learning and what is motor adaptation. Then, we will present a model of eye-hand co-ordination during self-moved target tracking, which we used as a way to specifically address these topics. Finally, we will show some evidence about the use of proprioceptive information for updating the internal models, in the context of eye-hand co-ordination. Motor and afferent information appears to contribute to the parametric adjustment (adaptation) between arm motor command and visual information about arm motion. The study reported here was aimed at assessing the contribution of arm proprioception in building (learning) and updating (adaptation) these representations. The subjects (including a deafferented subject) had to make back and forth movements with their forearm in the horizontal plane, over learned amplitude and at constant frequency, and to track an arm-driven target with their eyes. The dynamical conditions of arm movement were altered (unexpectedly or systematically) during the movement by changing the mechanical properties of the manipulandum. The results showed a significant change of the latency and the gain of the smooth pursuit system, before and after the perturbation for the control subjects, but not for the deafferented subject. Moreover, in control subjects, vibrations of the arm muscles prevented adaptation to the mechanical perturbation. These results suggest that in a self-moved target tracking task, the arm motor system shares with the smooth pursuit system an internal representation of the arm dynamical properties, and that arm proprioception is necessary to build this internal model. As suggested by Ghez et al. (1990) (Cold Spring Harbor Symp. Quant. Biol., 55: 837-8471), proprioception would allow control subjects to learn the inertial properties of the limb.

Force-field adaptation without proprioception: Can vision be used to model limb dynamics?

Because our environment and our body can change from time to time, the efficiency of human motor behavior relies on the updating of the neural processes transforming intentions into actions. Adaptation to the context critically depends on sensory feedback such as vision, touch or hearing. Although proprioception is not commonly listed as one of the main senses, its role is determinant for the coordination of daily gestures like goal-directed arm movements. In particular, previous work suggests that proprioceptive information is critical to update the internal representation of limb dynamic properties. Here, we examined the motor behavior of a deafferented patient, deprived of proprioception below the nose, to assess adaptation to new dynamic conditions in the absence of limb proprioception. The patient, and age-matched control participants, reached toward visual targets in a new force field created by a rotating platform. Full vision of the limb and workspace was available throughout the experiment. Although her impairment was obvious in baseline reaching performance, the proprioceptively deafferented patient clearly adapted to the new force conditions. In fact, her time course of adaptation was similar to that observed in controls. Moreover, when tested in the normal force field after adaptation to the new force field, the patient exhibited after-effects similar to those of controls. These findings show that motor adaptation to a modified force field is possible without proprioception and that vision can compensate for the permanent loss of proprioception to update the central representation of limb dynamics.

On-line versus off-line vestibular-evoked control of goal-directed arm movements.

The present study tested whether vestibular input can be processed on-line to control goal-directed arm movements towards memorized visual targets when the whole body is passively rotated during movement execution. Subjects succeeded in compensating for current body rotation by regulating ongoing arm movements. This performance was compared to the accuracy with which subjects reached for the target when the rotation occurred before the movement. Subjects were less accurate in updating the internal representation of visual space through vestibular signals than in monitoring on-line body orientation to control arm movement. These results demonstrate that vestibular signals contribute to motor control of voluntary arm movements and suggest that the processes underlying on-line regulation of goal-directed movements are different from those underlying navigation-like behaviors.

Visual feedback of the moving arm allows complete adaptation of pointing movements to centrifugal and Coriolis forces in human subjects

A classical visuo-manual adaptation protocol carried out on a rotating platform was used to test the ability of subjects to adapt to centrifugal and Coriolis forces when visual feedback of the arm is manipulated. Three main results emerge: (a) an early modification of the initial trajectory of the movements takes place even without visual feedback of the arm; (b) despite the change in the initial trajectory, the new external force decreases the accuracy of the pointing movements when vision is precluded; (c) a visual adaptive phase allows complete adaptation of the pointing movements performed in a modified gravitoinertial field. Therefore vision would be essential for subjects to completely adapt to centrifugal and Coriolis forces. However, other sensory signals (i.e. vestibular and proprioceptive) may constitute the basis for early but partial correction of the pointing movements.

Pointing to a target from an upright position in human: tuning of postural responses when there is target uncertainty

Human subjects performed, from a standing position, rapid hand pointings to visual targets located within or beyond the prehension space. To examine the interaction between posture and the goal-directed movement we introduced a visual double-step perturbation requiring a reprogramming of the hand movement. Trials directed towards the same spatial goal but differentiated only by the likeliness of a visual double-step were compared. The hand kinematics was not affected by the uncertainty of the visual perturbation; an increased trunk bending, however, was observed. This suggests that uncertainty constraints are integrated in a predictive manner for the optimal coordination of the hand and postural control systems.

Bayesian networks and information theory for audio-visual perception modeling

Thanks to their different senses, human observers acquire multiple information coming from their environment. Complex cross-modal interactions occur during this perceptual process. This article proposes a framework to analyze and model these interactions through a rigorous and systematic data-driven process. This requires considering the general relationships between the physical events or factors involved in the process, not only in quantitative terms, but also in term of the influence of one factor on another. We use tools from information theory and probabilistic reasoning to derive relationships between the random variables of interest, where the central notion is that of conditional independence. Using mutual information analysis to guide the model elicitation process, a probabilistic causal model encoded as a Bayesian network is obtained. We exemplify the method by using data collected in an audio-visual localization task for human subjects, and we show that it yields a well-motivated model with good predictive ability. The model elicitation process offers new prospects for the investigation of the cognitive mechanisms of multisensory perception.

Interaction between Reference Frames during Subjective Vertical Estimates in a Tilted Immersive Virtual Environment

Numerous studies highlighted the influence of a tilted visual frame on the perception of the visual vertical (‘rod-and-frame effect’ or RFE). Here, we investigated whether this influence can be modified in a virtual immersive environment (CAVE-like) by the structure of the visual scene and by the adjustment mode allowing visual or visuo-kinaesthetic control (V and VK mode, respectively). The way this influence might dynamically evolve throughout the adjustment was also investigated in two groups of subjects with the head unrestrained or restrained upright. RFE observed in the immersive environment was qualitatively comparable to that obtained in a real display (portable rod-and-frame test; Oltman 1968, Perceptual and Motor Skills 26 503–506). Moreover, RFE in the immersive environment appeared significantly influenced by the structure of the visual scene and by the adjustment mode: the more geometrical and meaningful 3-D features the visual scene contained, the greater the RFE. The RFE was also greater when the subjective vertical was assessed under visual control only, as compared to visuo-kinaesthetic control. Furthermore, the results showed a significant RFE increase throughout the adjustment, indicating that the influence of the visual scene upon subjective vertical might dynamically evolve over time. The latter effect was more pronounced for structured visual scenes and under visuo-kinaesthetic control. On the other hand, no difference was observed between the two groups of subjects having the head restrained or unrestrained. These results are discussed in terms of dynamic combination between coexisting reference frames for spatial orientation.

To transfer or not to transfer? Kinematics and laterality quotient predict interlimb transfer of motor learning

Humans can remarkably adapt their motor behavior to novel environmental conditions, yet it remains unclear which factors enable us to transfer what we have learned with one limb to the other. Here we tested the hypothesis that interlimb transfer of sensorimotor adaptation is determined by environmental conditions but also by individual characteristics. We specifically examined the adaptation of unconstrained reaching movements to a novel Coriolis, velocity-dependent force field. Right-handed subjects sat at the center of a rotating platform and performed forward reaching movements with the upper limb toward flashed visual targets in prerotation, per-rotation (i.e., adaptation), and postrotation tests. Here only the dominant arm was used during adaptation and interlimb transfer was assessed by comparing performance of the nondominant arm before and after dominant-arm adaptation. Vision and no-vision conditions did not significantly influence interlimb transfer of trajectory adaptation, which on average was significant but limited. We uncovered a substantial heterogeneity of interlimb transfer across subjects and found that interlimb transfer can be qualitatively and quantitatively predicted for each healthy young individual. A classifier showed that in our study, interlimb transfer could be predicted based on the subjects task performance, most notably motor variability during learning, and his or her laterality quotient. Positive correlations suggested that variability of motor performance and lateralization of arm movement control facilitate interlimb transfer. We further show that these individual characteristics can predict the presence and the magnitude of interlimb transfer of left-handers. Overall, this study suggests that individual characteristics shape the way the nervous system can generalize motor learning.

Accuracy of spatial localization depending on head posture in a perturbed gravitoinertial force field

Spatial orientation is crucial when subjects have to accurately reach memorized visual targets. In previous studies modified gravitoinertial force fields were used to affect the accuracy of pointing movements in complete darkness without visual feedback of the moving limb. Target mislocalization was put forward as one hypothesis to explain this decrease in accuracy of pointing movements. The aim of this study was to test this hypothesis by determining the accuracy of spatial localization of memorized visual targets in a perturbed gravitoinertial force field. As head orientation is involved in localization tasks and carrying relevant sensory systems (visual, vestibular and neck muscle proprioceptive), we also tested the effect of head posture on the accuracy of localization. Subjects (n=10) were seated off-axis on a rotating platform (120° s−1) in complete darkness with the head fixed (head-fixed session) or free to move (head-free session). They were required to report verbally the egocentric spatial localization of visual memorized targets. They gave the perceived target location in direction (i.e. left or right) and in amplitude (in centimeters) relative to the direction they thought to be straight ahead. Results showed that the accuracy of visual localization decreased when subjects were exposed to inertial forces. Moreover, subjects localized the memorized visual targets more to the right than their actual position, that was in the direction of the inertial forces. With further analysis, it appeared that this shift of localization was concomitant with a shift of the visual straight ahead (VSA) in the opposite direction. Thus, the modified gravitoinertial force field led to a modification in the orientation of the egocentric reference frame. Furthermore, this shift of localization increased when the head was free to move while the head was tilted in roll toward the center of rotation of the platform and turned in yaw in the same direction. It is concluded that the orientation of the egocentric reference frame was influenced by the gravitoinertial vector.