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Coriolis-force-induced trajectory and endpoint deviations in the reaching movements of labyrinthine-defective subjects.

When reaching movements are made during passive constant velocity body rotation, inertial Coriolis accelerations are generated that displace both movement paths and endpoints in their direction. These findings directly contradict equilibrium point theories of movement control. However, it has been argued that these movement errors relate to subjects sensing their body rotation through continuing vestibular activity and making corrective movements. In the present study, we evaluated the reaching movements of five labyrinthine-defective subjects (lacking both semicircular canal and otolith function) who cannot sense passive body rotation in the dark and five age-matched, normal control subjects. Each pointed 40 times in complete darkness to the location of a just extinguished visual target before, during, and after constant velocity rotation at 10 rpm in the center of a fully enclosed slow rotation room. All subjects, including the normal controls, always felt completely stationary when making their movements. During rotation, both groups initially showed large deviations of their movement paths and endpoints in the direction of the transient Coriolis forces generated by their movements. With additional per-rotation movements, both groups showed complete adaptation of movement curvature (restoration of straight-line reaches) during rotation. The labyrinthine-defective subjects, however, failed to regain fully accurate movement endpoints after 40 reaches, unlike the control subjects who did so within 11 reaches. Postrotation, both groups' movements initially had mirror image curvatures to their initial per-rotation reaches; the endpoint aftereffects were significantly different from prerotation baseline for the control subjects but not for the labyrinthine-defective subjects reflecting the smaller amount of endpoint adaptation they achieved during rotation. The labyrinthine-defective subjects' movements had significantly lower peak velocity, higher peak elevation, lower terminal velocity, and a more vertical touchdown than those of the control subjects. Thus the way their reaches terminated denied them the somatosensory contact cues necessary for full endpoint adaptation. These findings fully contradict equilibrium point theories of movement control. They emphasize the importance of contact cues in adaptive movement control and indicate that movement errors generated by Coriolis perturbations of limb movements reveal characteristics of motor planning and adaptation in both healthy and clinical populations.

Aged↗

A QUANTITATIVE, THREE-DIMENSIONAL METHOD FOR ANALYZING ROTATIONAL MOVEMENT FROM SINGLE-VIEW MOVIES

The study of animal movement is an important aspect of functional morphological research. The three-dimensional movements of (parts of) animals are usually recorded on two-dimensional film frames. For a quantitative analysis, the real movements should be reconstructed from their projections. If movements occur in one plane, their projection is distorted only if this plane is not parallel to the film plane. Provided that the parallel orientation of the movement with respect to the film plane is checked accurately, a two-dimensional method of analysis (ignoring projection errors) can be justified for quantitative analysis of planar movements. Films of movements of skeletal elements of the fish head have generally been analyzed with the two-dimensional method (e.g. Sibbing, 1982; Hoogenboezem et al. 1990; Westneat, 1990; Claes and de Vree, 1991), which is justifiable for planar movements. Unfortunately, the movements of the head bones of fish are often strongly non-planar, e.g. the movement of the pharyngeal jaws and the gill arches. The two-dimensional method is inappropriate for studying such complex movements (Sibbing, 1982; Hoogenboezem et al. 1990). For a qualitative description of movement patterns, the conditions for the use of the two-dimensional method may be somewhat relaxed. When two (or more) views of a movement are recorded simultaneously, the three-dimensional movements can readily be reconstructed using two two-dimensional images (e.g. Zarnack, 1972; Nachtigall, 1983; van Leeuwen, 1984; Drost and van den Boogaart, 1986). However, because of technical (and budget) limitations, simultaneous views of a movement cannot always be shot. In this paper, a method is presented for reconstructing the three-dimensional orientation and rotational movement of structures using single-view films and for calculating rotation in an object-bound frame. Ellington (1984) presented a similar method for determining three-dimensional wing movements from single-view films of flying insects. Ellington's method is based upon the bilateral symmetry of the wing movements. The present method does not depend on symmetry and can be applied to a variety of kinematic investigations. It eliminates a systematic error: the projection error. The measuring error is not discussed; it is the same in the two-dimensional and three-dimensional method of analysis.

Journal Article↗

Discharges of superior colliculus neurons during head and eye movements of the alert cat.

452 single neurons from the superior colliculus were recorded in awake and non-paralysed cats. 75 neurons were obtained from cats with unrestrained horizontal head movements. 228 neurons remained unaffected by saccadic eye movements. Eye movement related discharge followed the onset of saccades in 156 neurons either only in the presence of a visual pattern (92 neurons) or in darkness, too (64 neurons). The latter reaction type probably depends on eye muscle afferents. In 48 neurons eye movements related activity preceded the onset of eye movements. 12 neurons eye movement related activity preceded the onset of eye movements. 12 neurons fired in synchrony with eye movements of any direction (type I). 30 neurons were excited during contralaterally directed eye versions within or into the contralateral head related hemifield. They were inhibited when the eyes moved within or into the ipsilateral head related hemifield (type II). 6 neurons with constant maintained activity during fixation were inhibited by ipsilaterally directed saccades, but remained unaffected by contralateral eye movements. Head movement related discharge followed the onset of head movements in 20 neurons only in presence of a visual pattern and also in darkness in 6 neurons. Ipsilateral head movements or postures strongly suppressed maintained activity and visual responsiveness of some neurons. 15 neurons discharged in synchrony with and prior to contralateral head movements. Ipsilateral head movements inhibited these neurons. Activation or inhibition were usually related to movement and to posture, exceptionally to movement or to posture. Electrical stimulation of recording sites of these neurons through the recording microelectrode elicits contralateral head movements.

Animals↗

Dynamic interactions of eye and head movements when reading with single-vision and progressive lenses in a simulated computer-based environment.

PURPOSE: To assess dynamic interactions of eye and head movements during return-sweep saccades (RSS) when reading with single-vision (SVL) versus progressive-addition (PAL) lenses in a simulated computer-based business environment. METHODS: Horizontal eye and head movements were recorded objectively and simultaneously at a rate of 60 Hz during reading of single-page (SP; 14 degrees horizontal [H]) and double-page (DP; 37 degrees H) formats at 60 cm with binocular viewing. Subjects included 11 individuals with normal presbyopic vision aged 45 to 71 years selected by convenience sampling from a clinic population. Reading was performed with three types of spectacle lenses with a different clear near field of view (FOV): a SVL (60 degrees H clear FOV), a PAL-I with a relatively wide intermediate zone (7.85 mm; 18 degrees H clear FOV), and a PAL-II with a relatively narrow intermediate zone (5.60 mm; 13 degrees H clear FOV). RESULTS: Eye movements were initiated before head movements in the SP condition, and the reverse was found in the DP condition, with all three lens types. Duration of eye movements increased as the zone of clear vision decreased in the SP condition, and they were longer with the PALs than with the SVL in the DP condition. Gaze stabilization occurred later with the PALs than with the SVL in both the SP and DP conditions. The duration of head movements was longer with the PAL-II than with the SVL in both the SP and DP conditions. Eye movement peak velocity was greater with the SVL than the PALs in the DP condition. CONCLUSIONS: Eye movement and head movement strategies and timing were contingent on viewing conditions. The longer eye movement duration and gaze-stabilization times suggested that additional eye movements were needed to locate the clear-vision zone and commence reading after the RSS. Head movements with PALs for the SP condition were similarly optically induced. These eye movement and head movement results may contribute to the reduced reading rate and related symptoms reported by some PAL wearers. The dynamic interactions of eye movements and head movements during reading with the PALs appear to be a sensitive indicator of the effect of lens optical design parameters on overall reading performance, because the movements can discriminate between SVL and PAL designs and at times even between PALs.

Aged↗

Electromyographic responses to constant position errors imposed during voluntary elbow joint movement in human.

The role of reflexes in the control of stiffness during human elbow joint movement was investigated for a wide range of movement speeds (1.5-6 rad/s). The electromyographic (EMG) responses of the elbow joint muscles to step position errors (step amplitude 0.15 rad; rise time 100 ms) imposed at the onset of targeted flexion movements (1.0 rad amplitude) were recorded. For all speeds of movement, the step position disturbance produced large modulations of the usual triphasic EMG activity, both excitatory and inhibitory, with an onset latency of 25 ms. In the muscles stretched by the perturbation, the early EMG response (25-60 ms latency) magnitude was greater than 50% of the activity during the unperturbed movements (background activity). In all muscles the EMG responses integrated over the entire movement were greater than 25% of the background activity. The responses were relatively greater for slower movements. Perturbations assisting the movement caused a short-latency (25-60 ms) reflex response (in the antagonist muscle) that increased with movement speed and was constant as a percentage of the background EMG activity. In contrast, perturbations resisting the movement caused a reflex response (in the agonist muscle) that was of the same absolute magnitude at all movement speeds, and thus decreased with movement speed as a percentage of the background EMG activity. There was a directional asymmetry in the reflex response, which produced an asymmetry in the mechanical response during slow movements. When the step perturbation occurred in a direction assisting the flexion movement, the antagonist muscle activity increased, but the main component of this response was delayed until the normal time of onset of the antagonist burst. When the step perturbation resisted the movement the agonist muscles responded briskly at short latency (25 ms). A reflex reversal occurred in two of six subjects. A fixed reflex response occurred in the antagonist muscle, regardless of the perturbation direction. For the extension direction perturbations (resisting movement), this response represented a reflex reversal (50 ms onset latency) and it caused the torque resisting the imposed step (stiffness) to drop markedly (below zero for one subject). Reflex responses were larger when the subject was prevented from reaching the target. That is, when the perturbation remained on until after the normal time of reaching the target, the EMG activity increased, with a parallel increase in stiffness. Similarly, when the perturbations prevented the subject from reaching the target during a 1-rad voluntary cyclic movement, the EMG and stiffness increased markedly.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult↗

Evidence of a limited visuo-motor memory used in programming wrist movements.

Human subjects can pre-program movements on the basis of visual cues. Experience in a particular task leads to the storage of appropriate control parameters which are used in programming subsequent movements, via a short-term motor memory. The form, duration and usage of this memory are, however, uncertain. Repetitive wrist flexion and extension movements were measured in four subjects. Three were neurologically normal men; the fourth subject had a peripheral large-fibre sensory neuropathy, depriving him of proprioceptive information about wrist movement. Subjects made alternating 45 degrees wrist movements between two visual targets; visual feedback of wrist position was provided for the first part of each trial. After 10 s of tracking, the subjects paused for an interval of 0-24 s before resuming tracking without visual feedback of wrist position. The positional accuracy of subsequent movements was analysed with respect to pause interval. Movement accuracy was reduced by the removal of visual feedback in all four subjects: movements after the pause interval were less accurate than those before the pause. Errors also accumulated within each sequence of movements made without visual feedback. Analysis of the first movement in each trial after the pause indicated a clear relationship between movement accuracy and pause interval. In all four subjects, movement accuracy decayed with longer pause intervals. In the deafferented subject, manipulation of the visual inputs (requiring visual fixation, rather than normal pursuit of the target; or direct viewing of the hand instead of viewing a cursor on a computer screen) affected the relationship between pause interval and subsequent movement accuracy. We propose that the memory used when producing these movements is a short-lasting visuo-motor signal, lasting a few seconds, which is derived from visual knowledge of previous movements, rather than a memory of a particular motor output. This visuo-motor signal is used to scale the amplitude of subsequent wrist movements. The brevity of the visuo-motor memory and the resultant inaccuracy of this deafferented subject and of our neurologically normal subjects implies that human feedforward control of the amplitude and position of wrist movements is severely limited.

Adult↗

Velocity curves of human arm and speech movements.

The velocity curves of human arm and speech movements were examined as a function of amplitude and rate in both continuous and discrete movement tasks. Evidence for invariance under scalar transformation was assessed and a quantitative measure of the form of the curve was used to provide information on the implicit cost function in the production of voluntary movement. Arm, tongue and jaw movements were studied separately. The velocity curves of tongue and jaw movement were found to differ in form as a function of movement duration but were similar for movements of different amplitude. In contrast, the velocity curves for elbow movements were similar in form over differences in both amplitude and duration. Thus, the curves of arm movement, but not those of tongue or jaw movement, were geometrically equivalent in form. Measurements of the ratio of maximum to average velocity in arm movement were compared with the theoretical values calculated for a number of criterion functions. For continuous movements, the data corresponded best to values computed for the minimum energy criterion; for discrete movement, values were in the range of those predicted for the minimum jerk and best stiffness criteria. The source of a rate dependent asymmetry in the form of the velocity curve of speech movements was assessed in a control study in which subjects produced simple raising and lowering movements of the jaw without talking. The velocity curves of the non-speech control gesture were similar in form to those of jaw movement in speech. These data, in combination with similar findings for human jaw movement in mastication, suggest that the asymmetry is not a direct consequence of the requirements of the task. The biomechanics and neural control of the orofacial system may be possible sources of this effect.

Arm↗

Speed-accuracy trade-off in the performance of pointing movements in different directions in two-dimensional space.

Nine healthy subjects performed 2D pointing movements using a joystick that controlled a screen cursor. Continuous visual feedback was provided until movement completion. Three variables were systematically manipulated: (1) target distance, (2) target size and (3) target direction. A four-way factorial ANOVA was used to analyze the effects of these fixed factors and of the random factor of subject on several movement parameters. Movement time increased with increasing distance and decreasing target size and as predicted from Fitts' law. The target direction did not affect movement time. In contrast the direction, distance and size of the target significantly affected the movement time until the first zero crossing on the speed record reflecting the time to bring the arm into the vicinity of the target. Movements on the lateral axis of the horizontal plane (horizontal movements) resulted in a decrease in initial movement time compared to movements on the anterior axis of the horizontal plane (vertical movements). A significant effect of target distance and direction but not target size was observed for the magnitude of maximum acceleration, maximum speed and maximum deceleration. Horizontal movements had a larger maximum acceleration, speed and deceleration. Furthermore the maximum speed and deceleration occurred earlier in time for these horizontal movements. Finally the number of secondary peaks on the speed record increased with decreasing target size and was not affected by the target distance or target direction. In conclusion our results indicate that different movement parameters are affected by target distance, size and direction. The crucial distinction was between parameters affected by target size and direction. These parameters did not overlap. Target direction affects the first part of movement execution while target size affects the final part of movement execution. Thus a clear segmentation of movement execution in two phases is supported by these results. The implications of these results for theoretical models of speed-accuracy trade-off are discussed.

Acceleration↗

Neuromagnetic fields accompanying unilateral and bilateral voluntary movements: topography and analysis of cortical sources.

Movement-related magnetic fields (MRMFs) accompanying left and right unilateral and bilateral finger flexions were studied in 6 right-handed subjects. Six different MRMF components occurring prior to, and during both unilateral and bilateral movements are described: a slow pre-movement readiness field (RF, 1-0.5 sec prior to movement onset); a motor field (MF) starting shortly before EMG onset; 3 separate "movement-evoked" fields following EMG onset (MEFI at 100 msec; MEFII at 225 msec; and MEFIII at 320 msec); and a "post-movement" field (PMF) following the movement itself. The bilateral topography of the RF and MF for both unilateral and bilateral movements suggested bilateral generators for both conditions. Least-squares fitting of equivalent current dipole sources also indicated bilateral sources for MF prior to both unilateral and bilateral movements with significantly greater strength of contralateral sources in the case of unilateral movements. Differences in pre-movement field patterns for left versus right unilateral movements indicated possible cerebral dominance effects as well. A single current dipole in the contralateral sensorimotor cortex could account for the MEFI for unilateral movements and bilateral sensorimotor sources for bilateral movements. Other MRMF components following EMG onset indicated similar sources in sensorimotor cortex related to sensory feedback or internal monitoring of the movement. The results are discussed with respect to the possible generators active in sensorimotor cortex during unilateral and bilateral movement preparation and execution and their significance for the study of cortical organization of voluntary movement.

Brain Mapping↗

Directional selectivity in the responses of units in cat primary visual cortex to passive eye movement.

The responses of single units in the primary visual cortex (Area 17) of anaesthetized, paralysed cats, to passive movement of the ipsilateral eye were studied. Responses to passive eye movement were found in about one-third of the cortical units isolated. Appropriate control experiments excluded visual, auditory and cutaneous inputs as the source of the effective signal during passive eye movement. The magnitudes of the responses to a number (usually four) of radial directions of passive eye movement were estimated from sets of peristimulus time histograms "interleaved" in time. Units were defined as "radially selective" if the responses to movement along one radius (e.g. vertically upwards) exceeded that along at least one other orthogonal radius (e.g. horizontal-temporal). Of 60 units tested, 53 (88%) were "radially selective" according to this definition. Some of the "radially selective" units showed an additional type of specificity to passive eye movement: (a) Some units responded preferentially to movement along one of the arcs of passive eye movement which were tested (e.g. vertical movement above the equator of the orbit). These units we have called "arc selective". (b) Other units were sensitive to the direction of movement and preferred movement in a particular direction over more than one arc (e.g. horizontal movement towards the temporal side in both nasal and temporal halves of the orbit). These we have called "direction selective". Twenty-one "radially selective" units showed one of these additional properties, nine were arc selective and twelve were direction selective. The implications of these results for the understanding of the function of orbital proprioceptive signals in the cortex are discussed briefly. Responses to passive eye movement were found in all of layers II-VI in Area 17 and the implications of this for the understanding of the pathway by which orbital proprioceptive signals reach the primary visual cortex are discussed. The experiments have shown that many units in cat visual cortex respond to passive eye movement and that most of these units have some specificity for particular radial directions of movement while some have additional specific properties. We believe that these properties of radial, directional and arc sensitivity are likely to be important in understanding the function of the orbital proprioceptive signal which arises during eye movement and they are particularly interesting in relation to the findings of others that this proprioceptive signal appears to be concerned in the normal development of visual properties in the cortex and in the control of visually guided movement in adult cats.

Animals↗

Control processes underlying elbow flexion movements may be independent of kinematic and electromyographic patterns: experimental study and modelling.

Using a non-linear dynamic model based on the lambda version of the equilibrium-point hypothesis, we investigated the shape and duration of the control patterns underlying discrete elbow movements. The model incorporates neural control variables, time-, position- and velocity-dependent intrinsic muscle and reflex properties. Two control variables (R and C) specify a positional frame of reference for activation of flexor and extensor motoneurons. The variable R (reciprocal command) specifies the referent joint angle (R) at which the transition of net flexor to extensor active torque or vice versa can be observed during changes in the actual joint angle elicited by an external force. The variable C (coactivation command) surrounds the transition angle by an angular range in which flexor and extensor muscles may be simultaneously active (if C > 0) or silent (if C < or = 0). An additional, time-dimensional control variable (mu command) influences the dependency of the threshold of the stretch reflex on movement velocity. This control variable is responsible for the reflex damping. Changes in the R command result in shifts in the equilibrium state of the system, a dynamical process leading to electromyographic modifications and movement production. Commands C and mu provide movement stability and effective energy dissipation preventing oscillations at the end of movement. A comparison of empirical and model data was carried out. A monotonic ramp-shaped pattern of the R command can account for the empirical kinematic and electromyographic patterns of the fastest elbow flexion movements made with or without additional inertia, as well as of self-paced movements. The rate of the shifts used in simulation was different for the three types of movements but independent of movement distance (20-80 degrees). This implies that, for a given type of movement, the distance is encoded by the duration of shift in the equilibrium state. The model also reproduces the kinematic and electromyographic patterns of the fastest uncorrected movements opposed in random trials by a high load (80-90% of the maximal) generated by position feedback to a torque motor. The following perturbation effects were simulated: a substantial decrease in the arm displacement (from 60-70 degrees to 5-15 degrees) and movement duration (to about 100 ms) so that these movements ended near the peak velocity of those which were not perturbed; a prolongation of the first agonist electromyographic burst as long as the load was applied; the suppression of the antagonist burst during the dynamic and static phases of movements: the reappearance of the antagonist burst in response to unloading accompanied by a short-latency suppression of agonist activity. These kinematic and electromyographic features of both perturbed and non-perturbed movements were reproduced by using the same control patterns which elicited a monotonic shift in the equilibrium state of the system ending before the peak velocity of non-perturbed movements. Our results suggest that the neural control processes underlying the fastest unopposed changes in the arm position are completed long before the end of the movement and phasic electromyographic activity. Neither the timing nor the amplitude of electromyographic bursts are planned but rather they represent the long-lasting dynamic response of central, reflex and mechanical components of the system to a monotonic, short-duration shift in the system's equilibrium state.

Adult↗

Preserved simple and impaired compound movement after infarction in the territory of the superior cerebellar artery.

A patient with an infarct in the distribution of the right superior cerebellar artery was studied with regard to his ability to make simple movements (visually triggered, self-terminated ballistic wrist movements), and compound movements (reaching to a visual target and precision pinch of a seen object). Movements on the right side of the body alone were affected. Control movements were made by the normal left upper extremity. Wrist movement on the right side was normal in reaction time, direction, peak velocity, and end-point position control as compared to the left. By contrast, both reaching and pinching movements on the right were impaired. Reaching movements showed marked decomposition of the compound elbow-shoulder movement into seriatim simple movements made alternately at elbow and shoulder. Pinching movements were not made, and instead winkling movements (a movement of index alone) were substituted. These results are compared to similar results of controlled inactivation of the cerebellar dentate nucleus in monkeys. We conclude that one function of the cerebellum may be to combine elements in the movement repertoires of downstream movement generators. When that ability is lost, a strategy may be voluntarily adopted of using the preserved simple movements in place of the impaired compound movements.

Arteries↗

Movement-related potentials in Parkinson's disease. Presence and predictability of temporal and spatial cues.

Activity of the supplementary motor area may be inferred from movement-related potentials (MRPs) which are associated with the preparation and execution of voluntary, or internally determined movements. Supplementary motor area activity may be abnormal in Parkinson's disease since its major input from the basal ganglia is disrupted. Investigation of the abnormalities in supplementary motor area activity associated with movement deficits in Parkinson's disease may therefore reveal functions of the basal ganglia and the supplementary motor area. Movement-related potentials associated with sequential movements were investigated under various cueing conditions in Parkinson's disease subjects and age-matched controls. In controls, MRPs revealed involvement of the supplementary motor area in movements which can be internally determined (non-cued and externally cued, predictable movements, but not unpredictable movements). In Parkinson's disease, however, the supplementary motor area was only involved in movements which must be internally determined (non-cued movements, but not externally cued movements); therefore impaired internal control mechanisms, operating via the supplementary motor area, are bypassed when external cues are given. As a result, Parkinson's disease patients are more reliant on external cues and are unable to use predictive models to internally guide movement. Supplementary motor area involvement also relied on the predictability (in controls) or presence (in Parkinson's disease) of timing cues and not spatial cues, indicating a role of the supplementary motor area and basal ganglia in the temporal organizations of sequential movement rather than the programming of specific movements. For non-cued movements, abnormalities in MRPs for Parkinson's disease subjects consisted of delayed MRP onset and peak times, and prolonged cortical activity following movement. These observations led to a proposed model of the interaction between the basal ganglia and the supplementary motor area, involving the temporal organization of voluntary and internally determined sequential movements.

Aged↗

Acoustic-induced eye movements.

This article reports the influence of moving acoustic signals on eye movements (oculomotor functions) using 350 healthy test subjects. An acoustic test setting developed especially for this purpose has made possible the application of moving acoustic signals of varying frequency and form (square stimuli, sinusoidal stimuli, circular movements), as well as the examination of visual-acoustic and vestibular-acoustic interactions. It must be stressed that according to these investigations, both voluntary and involuntary eye movements can be demonstrated in response to moving acoustic signals. Involuntary eye movements, however, under th given experimental conditions, only are observed in about 20% of all test subjects and their frequency of occurrence is highest in young female subjects. They are most pronounced in darkness and when there is no fixation, and they can be coupled to varying degrees with the given acoustic signal. Voluntary and involuntary eye movements do not differ very much from one another. They consist mainly of a series of saccades that form staircase jerks. In both cases, no typical smooth movements or nystagmoid movement forms occur--which are predominant in tracking eye movements of visual stimuli. Occasionally, slight drifting movements can be demonstrated in the intersaccadic intervals. At a larger movement amplitude and increased movement frequency of the sound signal, pure opposite jerks finally appear, which follow up to a movement frequency of 1 Hz. The eye movements are influenced only slightly by a change in the amplitude of the sound movement (+/- 15 degrees to +/- 90 degrees). However, a close phase relationship between eye and sound movement exists. If instead of continuous acoustic signals (music), single burst signals are given at an increasing repetition rate, this results (at low repetition rates of bursts) in orienting reactions upon the individual signals, while at high signal repetition rates (above 0.5 Hz), continuous eye movements in the form of staircase jerks result. In investigating acoustically induced tracking eye movements, many new questions have arisen that previously have been open only to psychophysical studies.

Auditory Pathways↗

Relation between red nucleus discharge and movement parameters in trained macaque monkeys.

Correlation and regression analyses were performed on thirty-three of the magnocellular red nucleus cells described in the previous paper. We sought to test for reliable relations between the parameters of individual tracking movements and corresponding bursts of neural discharge. High correlations were found between the following burst and movement parameters: (i) burst latency versus movement latency; (ii) burst duration versus movement duration; (iii) burst frequency versus movement velocity and (iv) number of spikes in the burst versus movement amplitude. Cells were ranked according to the average of the duration, velocity and amplitude correlation coefficients. The top twenty cells had average correlation coefficients ranging from 0.69 to 0.88 for their preferred movement. These cases were judged most likely to reveal the control functions of the red nucleus, and the following points refer to this sample. Burst onset led movement onset by 118 +/- 23 ms, and burst offset led movement offset by 50 +/- 38 ms. Burst duration increased as the duration of the movement increased (r = 0.87 +/- 0.11). The duration of the burst was approximately equal to movement duration (slope of 0.99 +/- 0.16) plus a constant (72 +/- 34 ms) throughout a broad range. Average discharge rate during the burst increased with average movement velocity (r = 0.69 +/- 0.15). The slope of the relation was 0.36 +/- 0.21 (pulses/s)/(deg/s) of joint rotation. The regression lines had consistent upward offsets (56 +/- 15 pulses/s) that exceeded the spontaneous discharge rate (17 +/- 10 pulses/s). The number of spikes in the burst increased with movement amplitude independent of velocity (r = 0.72 +/- 0.11). The slope of the relation was 0.62 spikes/deg and the offset was 13 +/- 4 spikes. The preferred movement was co-ordinated hand in fifteen cases, digit in three, elbow in one and shoulder in one. When these cells were tested with an alternate movement, the failure rate (cases in which a burst did not accompany a movement) increased from 1.4 to 20%, and the correlation coefficients generally were low and lacked significance. Cells in the top twenty had directionally specific responses, low variance in lead time, large depths of modulation (41-118 pulses/s) and low failure rates. Cells that failed to show strong parametric correlations often had one or more of the former attributes. It appears that high parametric correlations with individual movements are particularly restrictive criteria of relatedness.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Putaminal activity for simple reactions or self-timed movements.

To examine the role of basal ganglia-cortical circuits in movement initiation, we trained monkeys to make the same arm movements in two ways-in immediate reaction to a randomly timed external cue (cued movements) and also following a variable delay without an explicit initiation signal (self-timed movements). The two movement types were interleaved and balanced in overall timing to allow a direct comparison of activity before and during the movement. Posterior putaminal neurons generally had phasic, movement-related discharges that were comparable for cued and self-timed movements. On cued movements, neuronal activity increased sharply following cue onset. However, for self-timed movements, there was a slow build-up in activity that preceded the phasic discharge. This slow build-up was time-locked to movement and restricted to a narrow time window hundreds of milliseconds before movement. The difference in premovement activity between cued and self-timed trials was present before the earliest cue-onset times and was not related to any differences in the overall time-to-move between the two types of trials. These features suggest that activity evolving in the basal ganglia-cortical circuitry may drive the initiation of movements by increasing until an activity threshold is exceeded. The activity may increase abruptly in response to an external cue or gradually when the timing of movements is determined by the animals themselves rather than an external cue. In this view, small changes in activity that occur in advance of the much larger perimovement neuronal activity may be an important determinant of when movement occurs. In support of this hypothesis, we found that even for cued movements, faster reaction times were associated with slightly higher levels of activity hundreds of milliseconds before movement.

Animals↗

Control of sequential movements: evidence for generalized motor programs.

The neuromotor processes underlying the control of rapid sequential limb movements were investigated. Subjects learned to pronate and supinate their forearms rapidly to four target locations in a specific spatio-temporal pattern under two movement-time conditions. The response sequence was first performed in a total movement time of 600 ms. Subjects were then told to produce the movement as quickly as possible while ignoring any timing pattern that they had previously learned. Electromyographic (EMG) signals were recorded from the biceps brachii and pronator teres muscles. Kinematic and EMG analyses were performed to investigate the temporal characteristics underlying the two movement-time conditions. When subjects produced the response as quickly as possible, average movement time to perform each reversal movement decreased while average peak velocity increased. Average total movement time was reduced by approximately 100 ms. Although movement time decreased, the proportion of total time to perform each movement of the sequence remained essentially invariant between movement-time conditions. Similar results were obtained for velocity. The time at which peak velocity was achieved occurred earlier in absolute time, although when normalized to the proportion of total movement time, the time to reach peak velocity was also invariant. Thus subjects proportionally compressed the entire movement sequence in time. The EMG analysis demonstrated that total EMG time decreased 89 ms on the average when subjects sped up the movement sequence. Thus average burst durations for both the biceps and pronator teres muscles decreased when movement speed increased. When burst durations were normalized to a proportion of total EMG time, the average proportion of time each muscle was active remained invariant. Therefore, the temporal pattern of activity for the biceps and pronator teres muscles were also proportionally compressed. The present experiment provided additional evidence for the structure of generalized motor programs consisting of invariant and variant features. Movement speed was considered a variant feature, which is specified each time the program is executed. Relative timing, the proportion of total time to produce each segment of the response, was considered to be an invariant feature and inherent in the structure of the motor program. Support for the invariance of relative timing was observed at both the kinematic and neuromuscular levels of analyses. Alternative models (9-11, 24) were found inadequate to account for the invariance of relative timing with the variation in movement time observed in the present experiment.

Electromyography↗

EEG arousals and awakenings in relation with periodic leg movements during sleep.

It is known that periodic leg movements are frequently accompanied by full awakenings or by signs of EEG arousals. The time relationship of these EEG arousals with leg movements varies from patient to patient. They may precede or follow leg movements or occur simultaneously. It is not clear whether these arousals trigger leg movements or, alternatively, whether both EEG arousals and leg movements are separate expressions of a common pathophysiological mechanism. We investigated the temporal relationship of five EEG arousals, such as alpha activity, K-complexes, spindles, K-alpha, K-spindle activities and awakenings, with leg movements in 10 periodic leg movement patients. These EEG arousals were considered to be associated with leg movements if they occurred 10 s before/after or simultaneously with the onset of right or left tibialis muscle EMG potentials. It was found that 49.19% of EEG arousals occurred before leg movements, 30.61% occurred simultaneously and 23.18% occurred just after leg movements. The number of EEG arousals was significantly higher in the 10 s preceding leg movement than simultaneously or in the 10 s following. Alpha activity was the phenomenon associated most frequently with leg movements, irrespective of its temporal organization and was significantly higher during the 10 s preceding movement. Spindle and K-spindle activities were significantly higher before leg movement, whereas K-complex activity was significantly more frequent during leg movements. The number of awakenings was significantly higher after leg movements than simultaneously. These results indicated that leg movements are not primary, but rather are a phenomenon associated with an underlying arousal disorder.

Adult↗