Positioning Errors in Simple Targeted Movements of the Forearm Realized in the Absence of Visual Control: Effects of Vibrational Stimulation of Antagonist Muscles

In 17 healthy subjects, we examined the characteristics of targeted movements of the forearm, flexion from the initial position of full extension taken as 0 deg to a 50 deg target angle in the elbow joint (flexor tests, FTs) and extension from the initial angle of 100 deg to the same target angle (extensor tests, ETs) with return to the initial positions. A standard movement (its trajectory corresponded to a simple trapezium) was performed under conditions of visual feedback (the value of the target angle and trajectory of the movement were visualized on the screen of a monitor); then, this movement should be reproduced by the subject (according to an acoustic signal) in the absence of visual control. Target-reaching test movements in the absence of visual feedback differed from the standard ones in a higher velocity. Blindfold reproduction of standard movements realized under kinesthetic control was accompanied in all subjects by noticeable positive systematic errors of targeted positioning (in the group, on average, 5.16 ± 0.55 and 4.83 ± 0.58 deg under FT and ET conditions, respectively). Vibrational stimulation of the muscles whose activity mainly provided the movement and positioning (m. biceps brachii in the FT cases and m. triceps brachii in the case of ETs) resulted in decreases of the errors of kinesthetic positioning; intragroup means of these errors were 2.55 ± 0.36 deg (FTs) and 2.26 ± 0.40 deg (ETs). The positioning errors demonstrated even greater decreases upon vibrational stimulation of the muscles, which were relatively inactive under conditions of the tests and underwent passive stretching in the course of the movements (m. triceps in FTs and m. biceps in ETs). Mean intragroup values of the errors in these cases were 0.46 ± 0.25 and 0.52 ± 0.31 deg, respectively. The nature of systematic positioning errors in the reproduction of targeted movements in the absence of visual control and the mechanisms underlying the influence of vibrational stimulation of the muscles involved in realization of these movements on the positioning errors under kinesthetic control are discussed.     

Positioning of the Human Forearm in Tracking Movements and Their Reproduction under Conditions of Limited Visual Control

In 14 healthy persons, we studied movements of the forearm with its positioning on a target level. A double trapezium was used as the command trajectory (flexion in the elbow joint from the state of full extension, 0°, with positioning on the level of 50 or 60° and further flexion to the 100° angle, and a similar reverse movement). We compared (i) tracking movements, when the subject tried to adequately reproduce the movement of the target along the command trajectory visualized on the monitor screen and obtained visual information about the performed movement (shifts of the second light point in time/joint angle coordinates), and (ii) reproduction of these movements under conditions of limitation of the visual feedback (when there was no information about the performed movement). Parameters of the tracking movements and of their reproductions (delays of initiation of the movement phases as compared with the command signal, durations of these phases, and angle velocities of the forearm movement), as well as the quality of positioning after oppositely directed movements, were compared. Positioning on the target level performed under proprioceptive control (when visual control was limited) was accompanied by systematic errors, whose sign in most test series performed by most subjects coincided with the direction of the preceding movement phase. The pattern of signs of systematic positioning errors after movements of opposite directions was quite individual (typical of a given subject) and demonstrated no dependence on the value of the extensor loading. Averaged intragroup systematic errors of positioning after movement phase 1 (flexion to the target level) and phase 3 (extension to the same level) under conditions of a minimum extensor loading (0.5-1.0 N · m) were 2.57° and 2.52°, respectively. When the loading was substantial (3.6-6.0 N · m), the respective errors were 3.85° and 3.48°. The nonlinear properties of muscle stretch receptors in the elbow flexors and extensors (responsible for the significant dependence of the parameters of afferent signals produced in these receptors on the movement prehistory) are considered the primary reason for systematic errors when positioning is performed exclusively under proprioceptive control. The influence of alpha-gamma co-activation in active muscles on the characteristics of the above signals is discussed.     

Movements of the Forearm and Shoulder Performed against the Gravitation Force: Target Positioning under Kinesthetic Control

In humans, we tested targeted movements of the forearm and shoulder performed in the vertical direction (in a parallel manner with respect to the sagittal plane). Movements were realized, first, with the possibility for visual control of the coincidence of the angle of the limb link axis vs the vertical and the target angle value (using an optic system and video recording), and, second, in the absence of the above control. Movements including flexion (i.e., movement against the gravitation force) – extension of the limb link with an individually selected convenient velocity were initiated and terminated according to the presentation of permissive sound signals; simultaneously, EMGs were recorded from a few muscles flexing and extending the elbow and shoulder joints. We analyzed systematic errors of target positioning of the forearm and shoulder in movements realized exclusively under kinesthetic control. In the case of isolated flexion of the forearm for a 90 deg target angle, such errors in all members of the examined group (n = 11) were positive. These errors were, on average, 8.1 ± 0.7 deg without loading and reached 11.2 ± 0.9 deg with introduction of a 10 to 30 N additional loading on the forearm. Isolated movements of the shoulder for a 70 deg target angle (performed without loading, with full extension of the forearm and its voluntary fixation) were accompanied by positive errors of 18.3 ± 1.1 deg, on average. Both the movements and positioning were performed due to changes in the levels of activity of the flexor muscles, with minimum involvement of the antagonists. The nonlinear properties of the receptor apparatus responsible for the formation of a kinesthetic estimate of the joint angle (first of all, of muscle spindles) are a fundamental reason for positive errors of target positioning of the limb links realized under kinesthetic control in the absence of the visual one.     

Reproduction of tracking movements and target positioning of the forearm in humans in the absence of visual control

In 17 healthy volunteers, we studied movements of the forearm, which included episodes of positioning on the target level. The trajectory of the non-ballistic (relatively slow) movement looked like a double trapezium (flexion of the elbow joint from the state of full extension, 0 deg, positioning on the 50 deg level, further flexion to the limit angle of 100 deg, and a similar reverse sequence). The command trajectory and the trajectory of the realized movement were visualized with movements of cursors on a monitor in time/joint angle coordinates. We compared parameters of the tracking movements (in the presence of visual feedback) and their blindfold reproduction (with the complete absence of visual control). It was found that blindfold reproduction movements differ from sample tracking movements and their reproduction with partial limitation of visual control [16] in higher peak velocities and shorter durations, i.e., a trend toward conversion of such movements into ballistic ones was observed. Under conditions of elimination of visual control, movements that led to positioning were mostly hypermetric, i.e., positioning was usually accompanied by positive systematic errors (whose sign coincided with the direction of the preceding movement phase). The mean intragroup value of the systematic error of the first positioning (after flexion to the target level) was +6.73 ± 1.15 deg, while the respective mean for the second positioning (after extension to the same level) was +4.00 ± 1.31 deg. The nonlinear properties of stretch receptors of muscles whose activity provides the formation of a proprioceptive estimate of the joint angle are considered the crucial reason for systematic errors of blindfold positioning.Neirofiziologiya/Neurophysiology, Vol. 36, Nos. 5/6, pp. 393–404, September–December, 2004.This revised version was published online in April 2005 with a corrected cover date and copyright year.     

Systematic positioning errors in human voluntary single-joint movements without visual guidance

Healthy subjects underwent analysis for positioning accuracy. The flexion-extension movements in the ankle and elbow joints performed without visual control were studied. The movements were produced by flexor muscles against small background loading (extensors were inactive). The subject was asked to memorize a certain target value of the joint angle reached under visual guidance in the phase of flexor contraction. After the flexion- (additional activation of the flexors) or extension-directed movement (relaxation of these muscles) the subject was asked to restore the target level from memory, without visual guidance. In the first case, when the target level was finally approached due to the flexor relaxation, a systematic overshoot of the target joint angle was observed. In the second case, when the target level was finally approached due to the additional activation of the flexors after their temporal relaxation, the positioning was much more accurate.Neirofiziologiya/Neurophysiology, Vol. 26, No. 2, pp. 91–98, March–April, 1994.     

Peculiarities of Activation of Muscles of the Shoulder Belt in Voluntary Two-Joint Movements of the Upper Limb

We studied coordination of central motor commands (СMCs) coming to muscles of the shoulder and shoulder belt in the course of single-joint and two-joint movements including flexion and extension of the elbow and shoulder joints. Characteristics of rectified and averaged EMGs recorded from a few muscles of the upper limb were considered correlates of the CMC parameters. Special attention was paid to coordination of CMCs coming to two-joint muscles that are able to function as common flexors (m. biceps brachii, caput breve, BBcb) and common extensors (m. triceps brachii, caput longum, TBcl) of the elbow and shoulder joints. Upper limb movements used in the tests included planar shifts of the arm from one spatial point to another resulting from either simultaneous changes in the angles of the shoulder and elbow joints or isolated sequential (two-stage) changes in these joint angles. As was found, shoulder muscles providing movements of the elbow with changes in the angle of the elbow joint, i.e., BBcb and TBcl, were also intensely involved in the performance of single-joint movements in the shoulder joint. The CMCs coming to two-joint muscles in the course of two-joint movements appeared, in the first approximation, as sums of the commands received by these muscles in the course of corresponding single-joint movements in the elbow and shoulder joints. Therefore, if we interpret the isolated forearm movement performed due to a change in the angle of the elbow joint as the main motor event, while the shoulder movement is considered the accessory one, we can conclude that realization of a two-joint movement of the upper-limb distal part is based on superposition of CMCs related to basic movements (main and accessory). Neirofiziologiya/Neurophysiology, Vol. 41, No. 1, pp. 48–56, January–February, 2009.     

Different programming modes of human saccadic eye movements as a function of stimulus eccentricity: Indications of a functional subdivision of the visual field

1. Voluntary saccadic eye movements were made toward flashes of light on the horizontal meridian, whose duration and distance from the point of fixation were varied; eye movements were measured using d.c.-electrooculography.—2. Targets within 10°–15° eccentricity are usually reached by one saccadic eye movement. When the eyes turn toward targets of more than 10°–15° eccentricity, the first saccadic eye movement falls short of the target by an angle usually not exceeding 10°. The presence of the image of the target off the fovea (visual error signal) subsequent to such an undershoot elicits, after a short interval, corrective saccades (usually one) which place the image of the target on the fovea. In the absence of a visual error signal, the probability of occurrence of corrective saccades is low, but it increases with greater target eccentricities. These observations suggest that there are different, eccentricity-dependent modes of programming saccadic eye movements.—3. Saccadic eye movements appear to be programmed in retinal coordinates. This conclusion is based on the observations that, irrespective of the initial position of the eyes in the orbit, a) there are different programming modes for eye movements to targets within and beyond 10°–15° from the fixation point, and b_ the maximum velocity of saccadic eye movements is always reached at 25° to 30° target eccentricity. —4. Distributions of latency and intersaccadic interval (ISI) are frequently multimodal, with a separation between modes of 30 to 40 msec. These observations suggest that saccadic eye movements are produced by mechanisms which, at a frequency of 30 Hz, process visual information. —5. Corrective saccades may occur after extremely short intervals (30 to 60 msec) regardless of whether or not a visual error signal is present; the eyes may not even come to a complete stop during these very short intersaccadic intervals. It is suggested that these corrective saccades are triggered by errors in the programming of the initial saccadic eye movements, and not by a visual error signal. —6. The exitence of different, eccentricity-dependent programming modes of saccadic eye movements, is further supported by anatomical, physiological, psychophysical, and neuropathological observations that suggest a dissociation of visual functions dependent on retinal eccentricity. Saccadic eye movements to targets more eccentric than 10°–15° appear to be executed by a mechanism involving the superior colliculus (perhaps independent of the visual cortex), whereas saccadic eye movements to less eccentric targets appear to depend on a mechanism involving the geniculo-cortical pathway (perhaps in collaboration with the superior colliculus).     

Modulation of the soleus muscle H reflex caused by ballistic voluntary arm movements in humans

In healthy humans, we studied the influence of conditioning voluntary arm movements on the H reflex induced by transcutaneous stimulation of the tibial nerve and recorded from the soleus muscle. We examined the effects of flexion and extension of the forearm, as well as of finger clenching performed with the maximum rate. Conditioning arm movements were self-induced or realized upon presentation of a visual signal (light flash). We found that the pattern of changes in the H reflex is determined by the position of the subject’s body in the course of tests. The ipsilateral arm flexion in the elbow joint in the standing position resulted in depression of the H reflex lasting about 100 msec from the beginning of the movement, while the effect observed in the lying position (on the couch with the feet hanging free in the air) looked like a facilitation of the reflex lasting about 100 to 200 msec. The direction and dynamics of modifications of the H reflex under conditions of the use of different conditioning movements (forearm flexions/extensions and finger clenching of the ipsilateral arm, as well as contralateral forearm flexions in the elbow joint) were rather similar. We also showed that the observed facilitation of the H reflex began earlier than the voluntary arm movement (40 to 50 msec prior to the beginning). We hypothesize that these conditioning influences result from the action of central motor commands and represent the factor related to anticipatory postural rearrangements. Such rearrangements are directed toward the maintenance of equilibrium of the body in the course of a future movement. These commands depend significantly on the spatial position of the subject’s body. Neirofiziologiya/Neurophysiology, Vol. 40, No. 2, pp. 147–154, March–April, 2008.     

A method for the chronic electromagnetic recording of eye and head movements in monkeys

A search-coil method for two-dimensional chronic registration of eye and head movements is described. The method is based on the analysis of the electromotive force induced by a magnetic field in a search coil. The output parameters of the original dual system were measured using both the standard search coil and that implanted into monkey's eye. The precision of eye movement recording was evaluated in real time. Standard deviation of spontaneous noise level for both channels was equal to 0.16 degrees (deg). The same parameters representing eye movement error during gaze fixation in the horizontal (in the range of -20/+20 deg) and vertical (in the range of -13/+13 deg) directions were equal to 0.27-0.38 and 0.23-0.31 deg, respectively. The obtained errors were comparable with the angular size of the peripheral target stimuli (0.20 deg), which had to be traced by an animal with saccadic movements.     

Learning the dynamics of reaching movements results in the modification of arm impedance and long-latency perturbation responses

 Some characteristics of arm movements that humans exhibit during learning the dynamics of reaching are consistent with a theoretical framework where training results in motor commands that are gradually modified to predict and compensate for novel forces that may act on the hand. As a first approximation, the motor control system behaves as an adapting controller that learns an internal model of the dynamics of the task. It approximates inverse dynamics and predicts motor commands that are appropriate for a desired limb trajectory. However, we had previously noted that subtle motion characteristics observed during changes in task dynamics challenged this simple model and raised the possibility that adaptation also involved sensory–motor feedback pathways. These pathways reacted to sensory feedback during the course of the movement. Here we hypothesize that adaptation to dynamics might also involve a modification of how the CNS responds to sensory feedback. We tested this through experiments that quantified how the motor system's response to errors during voluntary movements changed as it adapted to dynamics of a force field. We describe a nonlinear approach that approximates the impedance of the arm, i.e., force response as a function of arm displacement trajectory. We observe that after adaptation, the impedance function changes in a way that closely matches and counters the effect of the force field. This is particularly prominent in the long-latency (>100 ms) component of response to perturbations. Therefore, it appears that practice not only modifies the internal model with which the brain generates motor commands that initiate a movement, but also the internal model with which sensory feedback is integrated with the ongoing descending commands in order to respond to error during the movement. Received: 10 January 2001 / Accepted in revised form: 30 May 2001     

Quantization of human motions and learning of accurate movements

This paper presents a mathematical model for the learning of accurate human arm movements. Its main features are that the movement is the superposition of smooth submovements, the intrinsic deviation of arm movements is considered, visual and kinesthetic feedback are integrated in the motion control, and the movement duration and accuracy are optimized with practice. This model is consistent with the jerky arm movements of infants, and may explain how the adult motion behavior emerges from the infant behavior. Comparison with measurements of adult movements shows that the kinematics of accurate movements are well predicted by the model. Received: 15 May 1997 / Accepted 5 December 1997     

Tests of a linear model of visual-vestibular interaction using the technique of parameter estimation

The goal of this study was to test whether a superposition model of smooth-pursuit and vestibulo-ocular reflex (VOR) eye movements could account for the stability of gaze that subjects show as they view a stationary target, during head rotations at frequencies that correspond to natural movements. Horizontal smooth-pursuit and the VOR were tested using sinusoidal stimuli with frequencies in the range 1.0–3.5 Hz. During head rotation, subjects viewed a stationary target either directly or through an optical device that required eye movements to be approximately twice the amplitude of head movements in order to maintain foveal vision of the target. The gain of compensatory eye movements during viewing through the optical device was generally greater than during direct viewing or during attempted fixation of the remembered target location in darkness. This suggests that visual factors influence the response, even at high frequencies of head rotation. During viewing through the optical device, the gain of compensatory eye movements declined as a function of the frequency of head rotation (P < 0.001) but, at any particular frequency, there was no correlation with peak head velocity (P > 0.23), peak head acceleration (P > 0.22) or retinal slip speed (P > 0.22). The optimal values of parameters of smooth-pursuit and VOR components of a simple superposition model were estimated in the frequency domain, using the measured responses during head rotation, as each subject viewed the stationary target through the optical device. We then compared the model's prediction of smooth-pursuit gain and phase, at each frequency, with values obtained experimentally. Each subject's pursuit showed lower gain and greater phase lag than the model predicted. Smooth-pursuit performance did not improve significantly if the moving target was a 10 deg × 10 deg Amsler grid, or if sinusoidal oscillation of the target was superimposed on ramp motion. Further, subjects were still able to modulate the gain of compensatory eye movements during pseudo-random head perturbations, making improved predictor performance during visual-vestibular interactions unlikely. We conclude that the increase in gain of eye movements that compensate for head rotations when subjects view, rather than imagine, a stationary target cannot be adequately explained by superposition of VOR and smooth-pursuit signals. Instead, vision may affect VOR performance by determining the context of the behavior. Received: 16 June 1997 / Accepted: 5 December 1997     

Planning and execution of hand movements

The presented approach focuses on the attempt to specify strategies of visually organizing sequences of different hand movements and the resulting fine-control of movement close to the target, since in skilled activities our ability to sequence a number of separate movements each having different spatiotemporal characteristics is of central importance. Sequences of different moves have been analysed here. After a gross distance covering part of the move, small correction movements are performed close to the target to reduce the position error. The length of the required correction movements and the corresponding positioning time increase with target distance. In order to investigate motor control strategies two different tasks, with and without time pressure, have been designed. Time pressure forces the subjects to finish the previous move and to prepare the next move simultaneously. Absolute, constant and variable errors revealed that under time pressure the subject changes the control strategy by increasing the constant and reducing the variable error.     

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. Received: 23 June 1994 / Accepted in revised form: 3 February 1995     

The speed of mitochondrial movement is regulated by the cytoskeleton and myosin in Picea wilsonii pollen tubes

Strategic control of mitochondrial movements and cellular distribution is essential for correct cell function and survival. However, despite being a vital process, mitochondrial movement in plant cells is a poorly documented phenomenon. To investigate the roles of actin filaments and microtubules on mitochondrial movements, Picea wilsonii pollen tubes were treated with two microtubule-disrupting drugs, two actin-disrupting drugs and a myosin inhibitor. Following these treatments, mitochondrial movements were characterized by multiangle evanescent wave microscopy and laser-scanning confocal microscopy. The results showed that individual mitochondria underwent three classes of linear movement: high-speed movement (instantaneous velocities >5.0 μm/s), low-speed movement (instantaneous velocities <5.0 μm/s) and variable-speed movement (instantaneous velocities ranging from 0.16 to 10.35 μm/s). 10 nM latrunculin B induced fragmentation of actin filaments and completely inhibited mitochondrial vectorial movement. Jasplakinolide treatment induced a 28% reduction in chondriome motility, and dramatically inhibition of high-speed and variable-speed movements. Treatment with 2,3-butanedione 2-monoxime caused a 61% reduction of chondriome motility, and the complete inhibition of high-speed and low-speed movements. In contrast to actin-disrupting drugs, microtubule-disrupting drugs caused mild effects on mitochondrial movement. Taxol increased the speed of mitochondrial movement in cortical cytoplasm. Oryzalin induced curved mitochondrial trajectories with similar velocities as in the control pollen tubes. These results suggest that mitochondrial movement at low speeds in pollen tubes is driven by myosin, while high-speed and variable-speed movements are powered both by actin filament dynamics and myosin. In addition, microtubule dynamics has profound effects on mitochondrial velocity, trajectory and positioning via its role in directing the arrangement of actin filaments.     

Long-lasting inhibition of the soleus muscle H reflex related to realization of voluntary arm movements in humans

In healthy humans, we recorded the H reflex induced by transcutaneous stimulation of the tibial nerve (recording from the soleus muscle). In subjects in the lying position, we studied changes in the H reflex values after preceding voluntary arm movements realized with a maximum velocity after presentation of an acoustic signal. On the 200th to 300th msec after forearm flexion, long-lasting inhibition of the H reflex developed following a period of initial facilitation and reached the maximum, on average, 700 msec from the moment of the movement. Flexion of the contralateral upper limb in the elbow joint induced deeper inhibition than analogous movement of the ipsilateral arm. Long-lasting clear inhibition of the H reflex developed after arm flexion in the elbow joint but was slightly expressed after finger clenching. After inhibition reached the maximum, its time course was satisfactorily approximated by a logarithmic function of the time interval between the beginning of the conditioning voluntary movement and presentation of the test stimulus. Durations of inhibition calculated using a regression equation were equal to 6.6 sec and 8.5 sec after ipsilateral and contralateral elbow-joint flexions, respectively. Inhibition was not eliminated under conditions of tonic excitation of motoneurons of the tested muscle upon voluntary foot flexion. Long-lasting inhibition of the H reflex was also observed after electrical stimulation-induced flexions of the upper limb. The obtained data indicate that movements of the upper limb cause reflex long-lasting presynaptic inhibition of the soleus-muscle H reflex that can play a noticeable role in redistribution of the muscle tone during motor activity. Neirofiziologiya/Neurophysiology, Vol. 40, No. 3, pp. 221–227, May–June, 2008.     

Muscle activity in rapid multi-degree-of-freedom elbow movements: solutions from a musculoskeletal model

The activity of certain muscles that cross the elbow joint complex (EJC) are affected by forearm position and forearm movement during elbow flexion/extension. To investigate whether these changes are based on the musculoskeletal geometry of the joint, a three-dimensional musculotendinoskeletal computer model of the EJC was used to estimate individual muscle activity in multi-degree-of-freedom (df) rapid (ballistic) elbow movements. It is hypothesized that this model could reproduce the major features of elbow muscle activity during multi-df elbow movements using dynamic optimal control theory, given a minimum-time performance criterion. Results from the model are presented and verified with experimental kinematic and electromyographic data from movements that involved both one-df elbow flexion/extension and two-df flexion/extension with forearm pronation/supination. The model demonstrated how the activity of particular muscles is affected by both forearm position and movement, as measured in these experiments and as previously reported by others. These changes were most evident in the flexor muscles and least evident in the extensor muscles. The model also indicated that, for specific one- and two-df movements, activating a muscle that is antagonistic or noncontributory to the movement could reduce the movement time. The major features of muscle activity in multi-df elbow movements appear to be highly dependent on the joint's musculoskeletal geometry and are not strictly based on neural influences or neuroanatomical substrates. Received: 9 May 1997 / Accepted in revised form: 8 December 1998     

A method for measuring endpoint stiffness during multi-joint arm movements

Current methods for measuring stiffness during human arm movements are either limited to one-joint motions, or lead to systematic errors. The technique presented here enables a simple, accurate and unbiased measurement of endpoint stiffness during multi-joint movements. Using a computer-controlled mechanical interface, the hand is displaced relative to a prediction of the undisturbed trajectory. Stiffness is then computed as the ratio of restoring force to displacement amplitude. Because of the accuracy of the prediction (< 1 cm error after 200 ms) and the quality of the implementation, the movement is not disrupted by the perturbation. This technique requires only 13 as many trials to identify stiffness as the method of Gomi and Kawato (1997, Biological Cybernetics 76, 163-171) and may, therefore, be used to investigate the evolution of stiffness during motor adaptation.     

Influence of adaptation to horizontal body position on pointing errors and visual estimation of a target position in humans

The central program of a targeted movement includes a component intended for to compensate for the weight of the arm; this is why the accuracy of pointing to a memorized position of the visual target in darkness depends on orientation of the moving limb in relation to the vertical axis. Transition from the vertical to the horizontal body position is accompanied by a shift of the final hand position along the body axis towards the head. We studied how pointing errors and visual localization of the target are modified due to adaptation to the horizontal body position; targeted movements to a real target were repeatedly performed during the adaptation period. Three types of experiments were performed: a basic experiment, and two different experiments with adaptation realized under somewhat dissimilar conditions. In the course of the first adaptation experiment, subjects received no visual information on the hand’s position in space, and targeted movements of the arm to a luminous target could be corrected using proprioceptive information only. With such a paradigm, the accuracy of pointing to memorized visual targets showed no adaptation-related changes. In the second adaptation experiment, subjects were allowed to continuously view a marker (a light-emitting diode taped to the fingertip). After such adaptation practice, the accuracy of pointing movements to memorized targets increased: both constant and variational errors, as well as both components of constant error (i.e.,X andY errors) significantly dropped. Testing the accuracy of visual localization of the targets by visual/verbal adjustment, performed after this adaptation experiment, showed that the pattern of errors did not change compared with that in the basic experiment. Therefore, we can conclude that sensorimotor adaptation to the horizontal position develops much more successfully when the subject obtains visual information about the working point position; such adaptation is not related to modifications in the system of visual localization of the target.     

The representation of arm movements in postcentral and parietal cortex

Considerable experimental evidence supports the hypothesis that the neocortical processes underlying kinesthetic sensation form a hierarchical series of cells signalling increasingly complex patterns of movement of the body. However, this view has been criticized and the data lack quantitative verification under controlled conditions. These studies have also typically used one-dimensional (reciprocal) movements, even of multiple degree-of-freedom joints such as the wrist or shoulder, and have been restricted to passive movements. This latter limitation is particularly critical, since the response of many muscle receptors is affected by fusimotor activity while that of many articular receptors is sensitive to the level of muscle contractile activity. Both factors introduce significant kinesthetic ambiguity to the signals arising from these receptors during active movement. This ambiguity is evident in the discharge of primary somatosensory cortex proprioceptive cells. Studies in area 5 show that single cells signal shoulder joint movements in the form of broad directional tuning curves. The pattern of activity of the entire population encodes movement direction. The cells appear to encode spatial aspects of movement unambiguously, since their discharge is relatively insensitive to the changes in muscle activity required to produce the same movements under different load conditions. It is not yet certain whether the somesthetic activity in area 5 is a kinesthetic representation that is sequential to and hierarchically superior to that in SI, or whether it is a parallel representation with separate and distinct function.