Ankle joint movements are encoded by both cutaneous and muscle afferents in humans.

We analyzed the cutaneous encoding of two-dimensional movements by investigating the coding of movement velocity for differently oriented straight-line movements and the coding of complex trajectories describing cursive letters. The cutaneous feedback was then compared with that of the underlying muscle afferents previously recorded during the same "writing-like" movements. The unitary activity of 43 type II cutaneous afferents was recorded in the common peroneal nerve in healthy subjects during imposed ankle movements. These movements consisted first of ramp-and-hold movements imposed at two different and close velocities in seven directions and secondly of "writing-like" movements. In both cases, the responses were analyzed using the neuronal population vector model. The results show that movement velocity encoding depended on the direction of the ongoing movement. Discriminating between two velocities therefore involved processing the activity of afferent populations located in the various skin areas surrounding the moving joint, as shown by the statistically significant difference observed in the amplitude of the sum vectors. Secondly, "writing-like" movements induced cutaneous neuronal patterns of activity, which were reproducible and specific to each trajectory. Lastly, the "cutaneous neuronal trajectories," built by adding the sum vectors tip-to-tail, nearly matched both the movement trajectories and the "muscle neuronal trajectories," built from previously recorded muscle afferents. It was concluded that type II cutaneous and the underlying muscle afferents show similar encoding properties of two-dimensional movement parameters. This similarity is discussed in relation to a central gating process that would for instance increase the gain of cutaneous inputs when muscle information is altered by the fusimotor drive.

Perceptual integration of illusory and imagined kinesthetic images.

It is generally agreed that motor imagery involves kinesthetic sensations especially as far as first-person imagery is concerned. It was proposed to determine the extent to which motor imagery and vibration-induced illusory sensations of movement are integrated perceptually. Imagined and illusory hand movements were evoked both separately and in various combinations in 12 volunteers. After each trial, the participants were asked to draw the movement trajectory perceived. In all the subjects, propriomimetic vibration patterns applied to various wrist muscles induced spatially oriented or more complex illusory hand movements such as writing or drawing. Depending on the instructions, the subjects were also able to produce imagined hand movements in various directions and at two different velocities. When straight illusory and imagined movements were evoked simultaneously, all the subjects perceived a single movement trajectory, in which the direction and the velocity of the two ongoing sensations were exactly integrated. This perceptual integration also occurred in the case of more complex movements, such as writing and drawing, giving rise to the perception of original trajectories also combining the features of both motor images. Because these two kinesthetic images, the one intentionally and centrally induced and the other peripherally evoked, activate almost the same neural network including cortical sensory and motor areas, parietal regions, and the cerebellum, these results suggest that common processes may be involved in such a perceptual fusion. The nature of these common processes is discussed, and some fields of research in which these findings could potentially be applied are suggested.

Predicting any arm movement feedback to induce three-dimensional illusory movements in humans.

Our sense of body posture and movement is mainly mediated by densely packed populations of tiny mechanoreceptors present in the muscles. Signals triggered in muscle spindles by our own actions contribute crucially to our consciousness of positions and movements by continuously feeding and updating dynamic sensorimotor maps. Deciphering the coding rules whereby the nervous system integrates this proprioceptive information perceptually could help to elucidate the mechanisms underlying kinesthesia. The aim of the present study was to test the validity of a "propriomimetic method" of predicting the proprioceptive streams emitted by each of the muscles involved in two- (2D) and three-dimensional (3D) arm movements. This method was based on the functional properties of muscle spindle populations previously recorded microneurographically in behaving humans. Ia afferent patterns mimicking those evoked when the "arm-forearm" ensemble is drawing straight lines, graphic symbols, and complex 3D figures were calculated. These simulated patterns were then delivered to the main elbow and shoulder muscle tendons of motionless volunteers via a set of vibrators. Results show that the simulated proprioceptive patterns applied induced, in passive subjects, illusory 2D and 3D arm movements, the trajectories of which were very similar to the expected ones. These simulated patterns can therefore be said to be a substitute for the Ia proprioceptive feedback evoked by any human arm movement and this method can certainly be extended to other musculoskeletal ensembles. The illusory movements induced when these proprioceptive patterns are applied to muscle groups via sets of vibrators may provide useful tools for sensorimotor rehabilitation purposes.

A new vibrator to stimulate muscle proprioceptors in fMRI.

Studying cognitive brain functions by functional magnetic resonance imaging (fMRI) requires appropriate stimulation devices that do not interfere with the magnetic fields. Since the emergence of fMRI in the 90s, a number of stimulation devices have been developed for the visual and auditory modalities. Only few devices, however, have been developed for the somesthesic modality. Here, we present a vibration device for studying somesthesia that is compatible with high magnetic field environments and that can be used in fMRI machines. This device consists of a poly vinyl chloride (PVC) vibrator containing a wind turbine and of a pneumatic apparatus that controls 1-6 vibrators simultaneously. Just like classical electromagnetic vibrators, our device stimulates muscle mechanoreceptors (muscle spindles) and generates reliable illusions of movement. We provide the fMRI compatibility data (phantom test), the calibration curve (vibration frequency as a function of air flow), as well as the results of a kinesthetic test (perceived speed of the illusory movement as a function of vibration frequency). This device was used successfully in several brain imaging studies using both fMRI and magnetoencephalography.

Countering postural posteffects following prolonged exposure to whole-body vibration: a sensorimotor treatment.

Postural stability of bulldozer operators after a day of work is investigated. When operators are no longer exposed to whole-body vibration (WBV) generated by their vehicle, their sensorimotor coordination and body representation remain altered. A sensorimotor treatment based on a set of customized voluntary movements is tested to counter and prevent potential post-work accidents due to prolonged exposure to WBV. This treatment includes muscle stretching, joint rotations, and plantar pressures, all known to minimize the deleterious effects of prolonged exposure to mechanical vibrations. The postural stability of participants (drivers; N = 12) was assessed via the area of an ellipse computed from the X and Y displacements of the center-of-pressure (CoP) in the horizontal plane when they executed a simple balance task before driving, after driving, and after driving and having performed the sensorimotor treatment. An ancillary experiment is also reported in which a group of non-driver participants (N = 12) performed the same postural task three times during the same day but without exposure to WBV or the sensorimotor treatment. Prolonged exposure to WBV significantly increased postural instability in bulldozer drivers after they operated their vehicle compared to prior to their day of work. The sensorimotor treatment allowed postural stability to return to a level that was not significantly different from that before driving. The results reveal that (1) the postural system remains perturbed after prolonged exposure to WBV due to operating a bulldozer and (2) treatment immediately after driving provides a "sensorimotor recalibration" and a significant decrease in WBV-induced postural instability. If confirmed in different contexts, the postural re-stabilizing effect of the sensorimotor treatment would constitute a simple, rapid, inexpensive, and efficient means to prevent post-work accidents due to balance-related issues.

Changes in human muscle spindle sensitivity during a proprioceptive attention task.

The aim of the present study was to test whether fusimotor control of human muscle spindle sensitivity changed when attention was selectively directed to the recognition of an imposed two-dimensional movement in the form of a written symbol. The unitary activities of 32 muscle spindle afferents (26 Ia, 6 II) were recorded by microneurography at the level of the common peroneal nerve. The patterns of firing rate in response to passive movements of the ankle, forming different letters or numbers, were compared in two conditions: control and recognition. No visual cues were given in either condition, but subjects had to recognize and name the character in one condition compared with not paying attention in the control condition. The results showed that 58% of the tested Ia afferents presented modified responses to movements when these had to be recognized. Changes in Ia afferent responses included decreased depth of modulation, increased variability of discharge, and changes in spontaneous activity. Not all changes were evident in the same afferent. Furthermore, the percentage of correctly recognized movements amounted to 63% when changes were observed, but it was only 48% when the primary ending sensitivity was unaltered. The responses of group II afferents were only weakly changed or unchanged. It is suggested that the altered muscle spindle sensitivity is because of selective changes in fusimotor control, the consequence of which might be to feed the brain movement trajectory information that is more accurate.

Relationship between the velocity of illusory hand movement and strength of MEG signals in human primary motor cortex and left angular gyrus.

We studied the relationship between the velocity of movement illusion and the activity level of primary motor area (M1) and of the left angular gyrus (AG) in humans. To induce illusory movement perception, we applied co-vibration at different frequencies on tendons of antagonistic muscle groups. Since it is well established that the velocity of illusory movement is related to the difference in vibration frequency applied to two antagonistic muscles, we compared magnetoencephalography (MEG) signals recorded in two conditions of co-vibration: in the "fast illusion" condition a frequency difference of 80 Hz was applied on the tendons of the right wrist extensor and flexor muscle groups, whereas in the "slow illusion" condition a frequency difference of 40 Hz was applied on the same muscle groups. The dipole strength, reflecting the activity level of structures, was measured over M1 and the left AG in two different time-periods: 0-400 and 400-800 ms in each condition. Our results showed that the activity level of the AG was similar in both conditions whatever the time-period, whereas the activity level of M1 was higher in the "fast illusion" condition compared to the "slow illusion" condition from 400 ms after the vibration onset only. The data suggest that the two structures differently contributed to the perception of illusory movements. Our hypothesis is that M1 would be involved in the coding of cinematic parameters of the illusory movement but not the AG.

Cortical correlates of illusory hand movement perception in humans: a MEG study.

The present study aimed to investigate cortical activity associated with perception of illusory hand movements elicited by tendon vibration using magnetoencephalography (MEG) in humans. We compared MEG responses in two conditions of stimulation, "illusion" and "no illusion". In the "illusion" condition, covibration at different frequencies applied on the tendons of the right wrist flexor and extensor muscle groups evoked illusory movements of the hand. In the "no illusion" condition, covibration was delivered at the same frequency on both tendon groups and no movement was perceived. In both experimental conditions, equivalent current dipoles (ECD) were identified in each of four time windows: 0-200 ms, 200-400 ms, 400-600 ms and 600-800 ms. Our data showed similar activation in S1, superior parietal gyrus and supramarginal gyrus in both conditions, whereas the supplementary motor area, M1 and the left angular gyrus were found active in the "illusion" condition only. Our results confirmed the role of posterior parietal areas as well as motor areas in the arising of kinesthetic sensations. The hypothesis of an interaction between the angular gyrus and the primary motor area occurring about 400 ms after the beginning of the stimulation is discussed.

Transcranial magnetic stimulation of the sensorimotor cortex alters kinaesthesia.

Tendon vibration is known to evoke perception of illusory movements, together with motor responses in the muscles antagonistic to those vibrated. In the present study, we assessed the perceptual and motor effects of transcranial magnetic stimulation of the sensorimotor cortex during illusions of hand movements evoked by vibration of wrist muscle tendons. The results showed that transcranial magnetic stimulation could accelerate or decelerate the illusory movements, depending on the site and intensity of magnetic stimulation. Whenever transcranial magnetic stimulation decelerated illusory movements, motor responses decreased, whereas whenever it accelerated illusory movements, motor responses increased. We conclude that motor responses associated with movement illusions have a cortical stage, because they are affected by experimentally induced disruption of activity in intracortical networks.

Kinesthetic illusions attenuate experimental muscle pain, as do muscle and cutaneous stimulation.

In the present study, muscle pain was induced experimentally in healthy subjects by administrating hypertonic saline injections into the tibialis anterior (TA) muscle. We first aimed at comparing the analgesic effects of mechanical vibration applied to either cutaneous or muscle receptors of the TA or to both types simultaneously. Secondly, pain alleviation was compared in subjects in whom muscle tendon vibration evoked kinesthetic illusions of the ankle joint. Muscle tendon vibration, which primarily activated muscle receptors, reduced pain intensity by 30% (p<0.01). In addition, tangential skin vibration reduced pain intensity by 33% (p<0.01), primarily by activating cutaneous receptors. Concurrently stimulating both sensory channels induced stronger analgesic effects (-51%, p<0.01), as shown by the lower levels of electrodermal activity. The strongest analgesic effects of the vibration-induced muscle inputs occurred when illusory movements were perceived (-38%, p=0.01). The results suggest that both cutaneous and muscle sensory feedback reduce muscle pain, most likely via segmental and supraspinal processes. Further clinical trials are needed to investigate these new methods of muscle pain relief.

Motor and parietal cortical areas both underlie kinaesthesia.

Tendon vibration has long been known to evoke perception of illusory movements through activation of muscle spindle primary endings. Few studies, however, have dealt with the cortical processes resulting in these kinaesthetic illusions. We conceived an fMRI experiment to investigate the cortical structures taking part in these illusory perceptions. Since muscle spindle afferents project onto different cortical areas involved in motor control it was necessary to discriminate between activation related to sensory processes and activation related to perceptual processes. To this end, we designed and compared different conditions. In two illusion conditions, covibration at different frequencies of the tendons of the right wrist flexor and extensor muscle groups evoked perception of slow or fast illusory movements. In a no illusion condition, covibration at the same frequency of the tendons of these antagonist muscle groups did not evoke a sensation of movement. Results showed activation of most cortical areas involved in sensorimotor control in both illusion conditions. However, in most areas, activation tended to be larger when the movement perceived was faster. In the no illusion condition, motor and premotor areas were little or not activated. Specific contrasts showed that perception of an illusory movement was specifically related to activation in the left premotor, sensorimotor, and parietal cortices as well as in bilateral supplementary motor and cingulate motor areas. We conclude that activation in motor as well as in parietal areas is necessary for a kinaesthetic sensation to arise.

Cutaneous afferents from human plantar sole contribute to body posture awareness.

We investigated whether the tactile information from the main supporting areas of the foot are used by the brain for perceptual purposes, namely body posture awareness and body representation in space. We applied various patterns of tactile stimulation to one or both soles of unmoving and blindfolded subjects by a 60 micro-vibrator tactile matrix set in a force platform. The perceptual effects of the stimulation were assessed through a 3D joystick handled by the subjects. All subjects reported illusory perceptions of whole-body leaning. Both orientation and amplitude of these perceptions depended on the stimulation pattern. Additional kinesthetic illusions sometimes occurred along the longitudinal axis of the body. We conclude that foot sole input contributes to the coding and the spatial representation of body posture.

Differential contributions of vision, touch and muscle proprioception to the coding of hand movements.

To further elucidate the mechanisms underlying multisensory integration, this study examines the controversial issue of whether congruent inputs from three different sensory sources can enhance the perception of hand movement. Illusory sensations of clockwise rotations of the right hand were induced by either separately or simultaneously stimulating visual, tactile and muscle proprioceptive channels at various intensity levels. For this purpose, mechanical vibrations were applied to the pollicis longus muscle group in the subjects' wrists, and a textured disk was rotated under the palmar skin of the subjects' right hands while a background visual scene was projected onto the rotating disk. The elicited kinaesthetic illusions were copied by the subjects in real time and the EMG activity in the adductor and abductor wrist muscles was recorded. The results show that the velocity of the perceived movements and the amplitude of the corresponding motor responses were modulated by the nature and intensity of the stimulation. Combining two sensory modalities resulted in faster movement illusions, except for the case of visuo-tactile co-stimulation. When a third sensory input was added to the bimodal combinations, the perceptual responses increased only when a muscle proprioceptive stimulation was added to a visuo-tactile combination. Otherwise, trisensory stimulation did not override bimodal conditions that already included a muscle proprioceptive stimulation. We confirmed that vision or touch alone can encode the kinematic parameters of hand movement, as is known for muscle proprioception. When these three sensory modalities are available, they contribute unequally to kinaesthesia. In addition to muscle proprioception, the complementary kinaesthetic content of visual or tactile inputs may optimize the velocity estimation of an on-going movement, whereas the redundant kinaesthetic content of the visual and tactile inputs may rather enhance the latency of the perception.

Combined contribution of tactile and proprioceptive feedback to hand movement perception.

UNLABELLED: Here we investigated how the tactile modality is used along with muscle proprioception in hand movement perception, whether these two sensory inputs are centrally integrated and whether they work complementarily or concurrently. The illusory right hand rotations induced in eleven volunteers by a textured disk scrolling under their hand in two directions at three velocities and/or by mechanical vibration applied to their wrist muscles at three frequencies were compared. The kinesthetic illusions were copied by the subjects on-line with their left hand. RESULTS: 1) in all the subjects, tactile stimulation alone induced an illusory hand rotation in the opposite direction to that of the disk, and the velocity of the illusion increased non-linearly with the disk velocity: the highest gain (the illusion velocity to disk velocity ratio) occurred at the slowest disk rotation; 2) adding a consistent proprioceptive stimulus increased the perceptual effects, whereas adding a conflicting proprioceptive stimulus of increasing frequency gradually decreased the tactile illusions and reversed their initial direction; 3) under both consistent and conflicting conditions, only strong proprioceptive stimulation significantly affected the gain of the resulting illusions, whereas the largest gain always occurred at low tactile stimulation levels when the illusory movements were in the same direction as the tactile-induced illusion. Tactile information may equal or even override muscle proprioceptive information in the perception of relatively small, slow hand movements. These two somatosensory inputs may be integrated complementarily, depending on their respective relevance to the task of accurately perceiving one's own hand movements.

Fusimotor drive may adjust muscle spindle feedback to task requirements in humans.

The aim of the present study was to investigate whether the fusimotor control of muscle spindle sensitivity may depend on the movement parameter the task is focused on, either the velocity or the final position reached. The unitary activities of 18 muscle spindle afferents were recorded by microneurography at the common peroneal nerve. We compared in two situations the responses of muscle spindle afferents to ankle movements imposed while the subject was instructed not to pay attention to or to pay attention to the movement, both in the absence of visual cues. In the two situations, three ramp-and-hold movements were imposed in random order. In one situation, the three movements differed by their velocity and in the other by the final position reached. The task consisted in ranking the three movements according to the parameter under consideration (for example, slow, fast, and medium). The results showed that paying attention to movement velocity gave rise to a significant increase in the dynamic and static responses of muscle afferents. In contrast, focusing attention on the final position reached made the muscle spindle feedback better discriminate the different positions and depressed its capacity to discriminate movement velocities. Changes are interpreted as reflecting dynamic and static gamma activation, respectively. The present results support the view that the fusimotor drive depends on the parameter the task is focused on, so that the muscle afferent feedback is adjusted to the task requirements.

Where is your shoulder? Neural correlates of localizing others' body parts.

Neuropsychological studies, based on pointing to body parts paradigms, suggest that left posterior parietal lobe is involved in the visual processing of other persons' bodies. In addition, some patients have been found with mild deficit when dealing with abstract human representations but marked impairment with realistically represented bodies, suggesting that this processing could be modulated by the abstraction level of the body to be analyzed. These issues were examined in the present fMRI experiment, designed to evaluate the effects of visually processing human bodies of different abstraction levels on brain activity. The human specificity of the studied processes was assessed using whole-body representations of humans and of dogs, while the effects of the abstraction level of the representation were assessed using drawings, photographs, and videos. To assess the effect of species and stimulus complexity on BOLD signal, we performed a two-way ANOVA with factors species (human versus animal) and stimulus complexity (drawings, photographs and videos). When pointing to body parts irrespective of the stimulus complexity, we observed a positive effect of humans upon animals in the left angular gyrus (BA 39), as suggested by lesion studies. This effect was also present in midline cortical structures including mesial prefrontal, anterior cingulate and precuneal regions. When pointing to body parts irrespective of the species to be processed, we observed a positive effect of videos upon photographs and drawings in the right superior parietal lobule (BA 7), and bilaterally in the superior temporal sulcus, the supramarginal gyrus (BA 40) and the lateral extrastriate visual cortex (including the "extrastriate body area"). Taken together, these data suggest that, in comparison with other mammalians, the visual processing of other humans' bodies is associated with left angular gyrus activity, but also with midline structures commonly implicated in self-reference. They also suggest a role of the lateral extrastriate cortex in the processing of dynamic and biologically relevant body representations.

Postural changes after sustained neck muscle contraction in persons with a lower leg amputation.

Lower leg amputation generally induces asymmetrical weight-bearing, even after rehabilitation treatment is completed. This is detrimental to the amputees' long term quality of life. In particular, increasing strains on joint surfaces that receive additional weight load causes back and leg pain, premature wear and tear and arthritis. This pilot study was designed to determine whether subjects with lower leg amputation experience postural post-effects after muscle contraction, a phenomenon already observed in healthy subjects, and whether this could improve the weight-bearing on their prosthesis. Fifteen subjects with a unilateral lower leg amputation and 17 control subjects volunteered to participate in this study. Centre of pressure (CP) position was recorded during standing posture, under eyes closed and open conditions. Recordings were carried out before the subjects performed a 30-s voluntary isometric lateral neck muscle contraction, and again 1 and 4 min after the contraction. Postural post-effects characterized by CP shift, occurred in the medio-lateral plane in the majority of the amputated (7/15 eyes closed, 9/15 eyes open) and control (9/17 eyes closed, 11/17 eyes open) subjects after the contraction. Half of these subjects had a CP shift towards the side of the contraction and the other half towards the opposite side. In four amputated subjects tested 3 months apart, shift direction remained constant. These postural changes occurred without increase in CP velocity. Thus, a 30-s voluntary isometric contraction can change the standing posture of persons with lower leg amputation. The post-effects might result from the adaptation of the postural frame of reference to the proprioceptive messages associated with the isometric contraction.

Inducing any virtual two-dimensional movement in humans by applying muscle tendon vibration.

In humans, tendon vibration evokes illusory sensation of movement. We developed a model mimicking the muscle afferent patterns corresponding to any two-dimensional movement and checked its validity by inducing writing illusory movements through specific sets of muscle vibrators. Three kinds of illusory movements were compared. The first was induced by vibration patterns copying the responses of muscle spindle afferents previously recorded by microneurography during imposed ankle movements. The two others were generated by the model. Sixteen different vibratory patterns were applied to 20 motionless volunteers in the absence of vision. After each vibration sequence, the participants were asked to name the corresponding graphic symbol and then to reproduce the illusory movement perceived. Results showed that the afferent patterns generated by the model were very similar to those recorded microneurographically during actual ankle movements (r=0.82). The model was also very efficient for generating afferent response patterns at the wrist level, if the preferred sensory directions of the wrist muscle groups were first specified. Using recorded and modeled proprioceptive patterns to pilot sets of vibrators placed at the ankle or wrist levels evoked similar illusory movements, which were correctly identified by the participants in three quarters of the trials. Our proprioceptive model, based on neurosensory data recorded in behaving humans, should then be a useful tool in fields of research such as sensorimotor learning, rehabilitation, and virtual reality.

Proprioceptive feedback enhancement induced by vibratory stimulation in complex regional pain syndrome type I: an open comparative pilot study in 11 patients.

OBJECTIVE: Complex regional pain syndrome type I (CRPS-I) is now considered as a central nervous system disease with peripheral manifestations. CRPS-I may result from a mismatch between sensory input and motor output leading to a disorganization of motor programming in cortical structures. According to previous studies in the field of motor control, one efficient way to correct this mismatch could be a proprioceptive feedback enhancement. The goal of the present study was to determine whether vibratory stimulation by improving proprioceptive feedback may increase range of motion and minimize pain in patients with CRPS-I. METHODS: An open non-randomized study was conducted in 11 patients with CRPS-I of the hand and wrist. Conventional rehabilitation sessions were given for 10 weeks. During each session, patients in the intervention group (n=7) received vibratory stimulation of the affected region; the remaining 4 patients served as the controls. RESULTS: After 10 weeks, range-of-motion gains were about 30% larger and pain severity was about 50% lower in the intervention group than in the control group. A significant decrease in analgesic use occurred in the intervention group. DISCUSSION: Vibratory stimulation may significantly improve range of motion and pain in patients with CRPS-I, probably by reestablishing consonance between sensory input and motor output at cortical level. Prospective randomized studies in larger numbers of patients are needed. Cross-over designs or simulated vibratory stimulation should be used to minimize bias.

Cutaneous afferents provide a neuronal population vector that encodes the orientation of human ankle movements.

The aim of this study was to analyse the directional coding of two-dimensional limb movements by cutaneous afferents from skin areas covering a multidirectional joint, the ankle. The activity of 89 cutaneous afferents was recorded in the common peroneal nerve, and the mean discharge frequency of each unit was measured during the outward phase of ramp and hold movements imposed in 16 different directions. Forty-two afferents responded to the movements in the following decreasing order (SA2, n = 24/27; FA2, n = 13/17; FA1, n = 3/24; SA1, n = 2/21). All the units activated responded to a specific range of directions, defining their 'preferred sector', within which their response peaked in a given direction, their 'preferred direction'. Based on the distribution of the preferred directions, two populations of afferents, and hence two skin areas were defined: the anterior and the external lateral parts of the leg. As the directional tuning of each population was cosine shaped, the neuronal population vector model was applied and found to efficiently describe the movement direction encoded by cutaneous afferents, as it has been previously reported for muscle afferents. The responses of cutaneous afferents were then considered with respect to those of the afferents from the underlying muscles, which were previously investigated, and an almost perfect matching of directional sensitivity was observed. It is suggested that the common movement-encoding characteristics exhibited by cutaneous and muscle afferents, as early as the peripheral level, may facilitate the central co-processing of their feedbacks subserving kinaesthesia.