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Biomedical subjects

W A MacKay

Publications and source records attributed to W A MacKay.

At least 19 recordsLinked to original sources

EEG rhythms of the sensorimotor region during hand movements.

The aim of this study was to determine what motor behaviors or conditions were associated with an increased occurrence of beta activity in the sensorimotor region of human subjects. EEG recordings were obtained from 8 electrodes symmetrically arranged around C3, with 3 cm interelectrode spacing. The electrode montage allowed calculation of the Laplacian operator at two positions, C3r and C3c, overlying the hand area of the motor cortex and of the somatosensory cortex, respectively. A variety of tasks involving right-hand movements of different levels of complexity, attention and preparation were performed. The corresponding EEG power spectra were subsequently computed for frequencies between 7 and 50 Hz. Repetitive hand movements alone (either drawing circles or writing one's signature) did not result in significantly increased beta activity in the sensorimotor region compared to relaxed conditions. However, both motor preparations and focused attention, whether movements were performed or not, were associated with an increase of high frequency beta activity (30-50 Hz) in the sensorimotor region. Therefore, the facilitatory effect of attention and motor preparation and not the functional activation of the sensorimotor cortex by hand movements was associated with an increase in synchronized fast beta activity.

Adolescent

Visual responses to reward-related cues in inferior parietal lobule.

A sample of 263 neurones was recorded in area 7a of the parietal lobe, in a monkey performing a reach task to visual targets displayed on a touch-sensitive videomonitor. The task had been operantly conditioned on food or juice rewards, and 78 (30%) of the units showed activity changes linked in some way to the reward. For most of these cells, the response was to the approach of the trainer's hand with the food reward. This specific visual response was similar irrespective of the direction of approach. Six cells increased discharge as soon as the task was completed in apparent anticipation of the reward. Another two neurones responded to missing a reward: they fired vigorously if the videoscreen was blanked in mid-trial because a target was not correctly touched. In many cases (40/78) the same cells responding to some aspect of the reward also responded to visual cues given during the task, especially the presentation of the target location. Reward-related activity in area 7a probably results from an integration of the visual and limbic inputs to this region, such that visual information which foretells behaviourally important events is emphasized.

Animals

Properties of reach-related neuronal activity in cortical area 7A.

1. In protocol 1, two macaque monkeys were trained to reach to illuminated buttons with the right arm as reach-related unit activity was monitored in area 7a of the left hemisphere. 2. Of 402 neurons recorded in area 7a, 109 changed their discharge rates during the reach task. The change could occur early or late in the trajectory, or during the return movement of the arm to the rest plate. Spatial preferences were seen in 59/109 reach-related cells, usually for the right or center buttons. 3. In protocol 2, another monkey was trained to reach with either arm to targets displayed on a touch-sensitive video monitor. Of 273 neurons sampled in area 7a (both hemispheres) during the bilateral task performance, 84 were reach-related: 33 responded similarly to reaches of either arm. Most of the rest had a contralateral arm preference. When bilateral reach-related cells had a spatial preference, that preference was the same for both arms. 4. With the use of two target sequences in either protocol, it was found that spatial preferences were observable only for primary reaches from the side of the body up to the target. Relatively few cells responded to other trajectories, and those that did usually failed to discriminate movement direction. Movement extent did not influence discharge rates. 5. Although a total of 125/270 reach cells had observable visual responses, only 4 out of 18 cells tested in both dark and light conditions showed a significant drop in reach-related activity in the dark. Thus visual input from the moving hand probably is responsible for only part of the reach activity in area 7a. 6. Reach-related activity in area 7a appears to signal specific phases of the motor performance and is often restricted to distinct spatial regions. As such, it could be used by the frontal lobe to facilitate upcoming elements of a motor sequence, including terminal corrections.

Animals

CNV, stretch reflex and reaction time correlates of preparation for movement direction and force.

Ten human subjects were tested in a pre-cued choice reaction time (RT) paradigm in which the warning stimulus gave a varying amount of prior information regarding the direction (flexion or extension) or force level (weak or strong) for an impending right forearm movement. During the preparatory period (PP), either CNV was monitored from 8 scalp leads, or elbow stretch reflexes were tested at selected times using mechanical torque steps as stimuli. Mean RTs increased as the amount of prior information decreased. The locus of maximal rate of increase of scalp negativity migrated from the frontal lobe to the parietal lobe during the PP, under all conditions. Using laplacian derivations, it was found that the CNV at Cz did not distinguish among the different information conditions, whereas the CNV over the somatosensory arm area was greatest when direction information was given in advance. The CNV over the arm motor region was greatest when force information was available. The last 100 msec of the PP was characterized by the development of current sources over the premotor region for full information, and over the somatosensory region for the other conditions. This coincided with the appearance of very large late stretch reflexes or triggered reactions in the prepared agonist for full information, indicating that the intended movement had been fully processed by this time and awaited only a sensory trigger. The data support a parametric model of motor preparation, with direction and force being processed by at least partially independent networks.

Adolescent

Reaching in the dark: enhanced responses in posterior parietal cortex.

In order to assess the relative importance of visual input to area 7 reach-related neuronal activity, a monkey was trained to reach to visual targets displayed on a video-monitor, both with and without visual feedback. Visual feedback was removed by having the monkey reach in darkness to a previously illuminated target. Of 19 reach-related cells recorded in area 7 both in the light and the dark, ten showed an enhancement of discharge in the dark. These included area 7b cells sensitive to screen contact and area 7a cells active during reach. Dark enhancement of active somatic responsiveness may partially compensate for the loss of visual guidance.

Animals

Changes in contingent-negative variation and reaction time related to precueing of direction and force of a forearm movement.

Scalp-recorded contingent-negative variation was analyzed in a reaction time paradigm with full, partial, or no prior information regarding two dimensions of a forearm response: direction (flexion/extension) and force level (weak/strong). Visual cues (light-emitting diodes) were used for the warning and response signals. The reaction time was shorter when direction, rather than force, was known in advance. Source derivation techniques revealed that the somatosensory arm area was more 'activated' by direction than force information, whereas the precentral cortex seemed to be more strongly influenced by force information. Partial advance information was sufficient to trigger preparatory activities specific for the revealed dimension of the ensuing movement.

Adult

Unit activity in the cerebellar nuclei related to arm reaching movements.

Single units in the fastigial, interpositus and dentate nuclei of two stump-tail macaque monkeys were studied in relation to a right arm, visually guided reaching task. Of 638 recorded cells, 149 showed activity changes correlated to the task, including 24 in the contralateral fastigial and interpositus. Reach-related discharge patterns fell into two broad categories, tonic and phasic. Tonic responses were maintained throughout the reach with no observable relation to kinematic parameters. Most of the task-related activity occurred during the upward lift of the arm toward the target button, with a drop-off as the arm was lowered toward the rest plate. Phasic response cells fired bursts (or suppressed discharge) at specific points in the arm trajectory, most commonly during the lift phase. Many had a sharp drop in discharge when the shoulder flexion torque was transiently reversed to decelerate the arm. For either type, restricted directional specificity was rarely seen in any nucleus, and correlations with recorded EMGs were weak. Visual responses to target button illumination were observed in both the fastigial and dentate nuclei, but did not necessarily correspond with the button giving the best movement-related response. Task-related activity changes started earliest in the fastigial nuclei and latest in the interpositus nuclei. The data suggested that cerebellar output facilitates motor centers in a rather general manner, but at precisely determined times.

Animals

Cerebellar nuclear activity in relation to simple movements.

Single unit activity in the fastigial, interpositus and dentate cerebellar nuclei was recorded in relation to simple elbow flexion and extension movements in two macaque monkeys. In common with proximal muscle activity, 94% of the task-related neurons had qualitatively similar discharge patterns for both directions of forearm movement. In many cases the flexion and extension discharge was virtually identical, but some cells had a distinct directional bias. The very few neurons which were directionally specific were located in the dentate and interpositus. Two had tonic activity well correlated to elbow angle. Task-related changes in discharge rate occurred earliest in dentate and latest in fastigial, but almost always during the period of concomitant proximal and elbow EMG changes. Correlations of phasic activity with movement velocity were uniformly weak. Many eye movement-related neurons were encountered in the fastigial, dentate and y-group nuclei. Fastigial eye cells, both bursting and tonic, tended to be highly direction specific, whereas dentate eye cells were usually omnidirectional and variable. For both arm and eye cerebellar cells, the directional preferences of phasic and tonic discharge, in the same neuron, could be opposed to one another.

Action Potentials

Neuronal correlates in posterior parietal lobe of the expectation of events.

During studies of response properties of single units in the posterior parietal cortex of 6 awake monkeys, 168 neurons were encountered (7.1% of examined units) which showed anticipatory types of activity. These neurons were found on either side of the intraparietal sulcus. In area 5, this expectation activity was expressed as a change in discharge rate whenever a specific body part (e.g. hand or shoulder) was approached by the investigator as though contact would be made. Invariably the neurons also responded to cutaneous and/or proprioceptive stimulation of the target body area. In area 7a the same type of response was also found but not always with a corresponding somatosensory receptive field. In addition, many neurons increased discharge rates (or rarely, decreased them) immediately prior to the expected occurrence of a reward, a visual task cue, or on hearing the approaching footsteps of a familiar person. None of these responses were correlated to eye movements, nor could they be attributed to any other body movement.

Action Potentials

Evoked potentials from passive elbow movements. II. Modification by motor intent.

Subjects were instructed to remain passive or to react to a forearm perturbation by opposing the imposed movement. Evoked potentials (EPs) were recorded at 8 scalp sites in both conditions. In the React condition, reflexes were observed in the EMG (mean onsets of 67 and 81 msec) and the EP was modified. Source derivation techniques revealed that the earliest cortical response (31 msec) across the central sulcus was not changed. Therefore the intention to react did not seem to affect afferent transmission to the primary sensorimotor cortex. Two periods of modulation were observed. In both, parietal and frontal potentials were modulated together, prior to the reflex components. After 70 msec, the pattern of potential gradients which occurred in the Passive case was accelerated and intensified in the React condition. The overall effect was to focus a larger zone of negativity over motor cortex at the time of triggered EMG output (109 msec). The earlier changes in cortical activity could be causally related to the appearance of the late stretch reflex. Since parietal and frontal areas were principally involved and not the motor area, it is suggested that the former exert a modulatory influence on spinal and brain-stem reflex centres.

Elbow

Measurements of human forearm viscoelasticity.

In human subjects, stiffness of the relaxed elbow was measured by three methods, using a forearm manipulandum coupled to a.d.c. torque motor. Elbow stiffness calculated from frequency response characteristics increased as the driving amplitude decreased. Step displacements of the forearm produced restoring torques linearly related to the displacement. The stiffness was very similar to that calculated from natural frequencies at amplitudes above 0.1 rad. Thirdly, elbow stiffness was estimated from brief test pulses, 120 ms in duration, by mathematically simulating the torque-displacement functions. Stiffness values in the limited linear range (under +/- 0.1 rad) were higher than in the linear range of the first two methods. A major component of elbow stiffness appears to decay within 1 s. The coefficients of viscosity determined from the simulation were, however, very similar to those calculated from the frequency response. Test pulse simulation was then used to determine joint impedance for different, actively maintained elbow angles. Joint stiffness and viscosity increased with progressive elbow flexion.

Adult

Properties of visual cue responses in primate precentral cortex.

Monkeys were trained to perform a visuomotor task involving the alignment of a cursor over a vertical target line on a videomonitor by flexion or extension movements of the wrist. The forelimb area of the contralateral precentral cortex was thoroughly explored during the task. Intracortical microstimulation was employed to classify the forelimb region into wrist flexion--extension and non-wrist flexion--extension populations. Unit recording revealed an initial response to the cue for movement, viz. the appearance of the cursor and target line on the videomonitor, while visual signals not related to the task failed to evoke any response. The mean latency of these visual cue responses was approximately 150 ms. A great majority of the responses (96%) were bidirectional in character, in that they did not correlate with the directional information embedded in the visual cue, nor were they good predictors for the direction or timing of the subsequent movement. They were uniformly distributed in both the wrist and non-wrist regions of the forelimb area; the non-forelimb areas were devoid of the cue response. Further, when the variability of response to the visual cue for the wrist and non-wrist populations was compared, no significant difference was observed. These observations are consistent with an interpretation that the visually triggered cue responses provide a generalized activation over the task-related area of precentral cortex, paving the way for later and more specific activations leading to the execution of the task.

Animals

Evoked potentials from passive elbow movements. I. Quantitative spatial and temporal analysis.

Evoked potentials following perturbations of the forearm were recorded monopolarly at 8 scalp sites. Successive 10 msec bins of individual EPs were compared across subjects to determine intervals of consistent potential change. From this analysis it was possible to objectively sort subjects into two subpopulations with different common wave forms. Spatial vectors (potential gradient between two adjacent leads) were computed as well as Laplacian derivatives, to identify electrodes closest to source activity. Subsequently, temporal derivatives of the vectors were computed to define the timing of statistically significant response phases. To more precisely localize the largest potential gradients, component vectors (or their derivatives) in the Laplacians were resolved algebraically on a scaled representation of the scalp. Convergence of these resultant vectors from adjacent areas, identified zones of significant potential change which corresponded to known somatosensory areas. The two subpopulations had initial responses of similar topography localized to the central sulcal region. Subsequently, for one group the area of activated cortex expanded to include posterior parietal and more frontal areas. Prestimulus negative potential shifts had different distributions for the two groups and are described in relation to the poststimulus differences.

Adult

Resonance properties of the human elbow.

In normal human subjects, elbow resonance was measured by driving the forearm over a range of frequencies. The relaxed forearm, in a manipulandum with a low inertial moment, had a resonance of 0.068 +/- 0.06 Hz which was remarkably consistent for either sex and over a wide variety of arm sizes. Elbow stiffness, therefore, appears to be well adapted to the forearm moment of inertia. The resonant frequency increased as the elbow muscles were cocontracted, but the amplitude of resonance dropped exponentially because of increased damping. By 25% of maximum electromyogram levels the elbow was critically damped. Comfortable voluntary motion exhibited an amplitude-period relation very similar to that of forearm resonance, suggesting that significant use was made of resonance properties. It was found that the forearm was voluntarily oscillated at its resonant frequency within a specific range, characteristic for the individual. Depending on movement frequency, resonance can provide anything from a major to a negligible contribution to voluntary oscillatory motion. It would greatly increase efficiency of performance when antagonist muscles are kept as relaxed as possible to minimize damping.

Adult

Stretch reflex modulation during a cyclic elbow movement.

Small torque pulses were delivered to the forearm in order to test the stretch reflex of the brachialis and triceps arm muscles in 11 normal subjects performing a cyclic movement about the elbow in the horizontal plane. The flexion-extension movement was paced by a metronome and performed under various loading conditions. Reflexes for each muscle were tested either in each 50 msec segment of the 2 sec cycle period, or in a smaller number of selected phases. A late reflex, appearing at a latency of about 60 msec (measured from the onset of the torque increment), was modulated extensively during the movement cycle. The amplitude of the late reflex increased markedly at the onset of a muscle contraction. In many of the subjects reflex responsiveness began to increase as early as 200 msec prior to the onset of voluntary muscle activity. Peak reflex responses were elicited by stimuli delivered 100-150 msec prior to the peak rate of increase of dynamic load (composed of inertial, viscous and elastic forces). The increase in responsiveness was followed by a drop which was generally coincident in time with the peak rate of increase of the load opposing muscle contraction. The modulation of the late reflex is appropriately timed for reflex-generated tension to help counteract dynamic loads, intrinsic to the movement.

Adult

Activity of primate precentral neurons during voluntary movements triggered by visual signals.

Awake, intact monkeys were trained to perform discrete flexion or extension movements of the hand about the wrist in response to visual signals. The object of the movement was to align a cursor, coupled to a manipulandum, on a target line. Cursor and target lines are displayed on a video monitor placed in front of the monkey. The target line was stepped to the right or left, randomly with regard to direction and timing, with each step implying an instruction for the monkey to make a voluntary movement for alignment. Single unit recording was made in the forelimb area of contralateral precentral cortex. Neurons were classified by their responses to passive sensory stimulation and the effects of local intracortical microstimulation into two populations; wrist flexion-extension (F-E) neurons, and all other forelimb neurons (non-wrist (F-E)). A significantly higher proportion of wrist (F-E) neurons as compared to non-wrist (F-E) neurons were task-related. Moreover the wrist (F-E) neurons exhibited exclusively reciprocal responses to the oppositely directed visual signals, whereas the non-wrist (F-E) neurons showed both reciprocal and bidirectional responses. No significant differences in mean latencies of responses, either in respect to the visual signals or to movement onset, were observed between the two populations of neurons. However the range of latencies in both instances was greater in the non-wrist (F-E) populations. The wrist (F-E) population showed significantly less response variability than the non-wrist (F-E) population with regard to response latencies to visual signals and movement onsets, and the degree of correlation between duration of response and reaction time.

Animals

Distribution of responses to visual cues for movement in precentral cortex or awake primates.

Unit recordings were made from areas 4 and 6 monkeys after they were trained to align a cursor over a vertical target line on a video screen by control of a manipulandum with wrist flexion or extension movement. The appearance of the cursor and line on the screen was the visual cue for movement. Responses were observed 150 (+/- 40) msec after cue presentation. The responses were found only in the forelimb area of precentral cortex, which was most immediately involved in the control of the task, and the majority of them were uncorrelated with either the specific details of the visual cue, or with the direction of the subsequent wrist movement.

Animals