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Biomedical subjects

C Ghez

Publications and source records attributed to C Ghez.

At least 55 records · Page 3Linked to original sources
PubMed1987

Trajectory control in targeted force impulses. I. Role of opposing muscles.

The functional role of opposing muscles in the production of isometric force trajectories was studied in six adult subjects producing impulses and steps of elbow flexor force, with different rise times and amplitudes. Rapidly rising forces were invariably associated with an alternating pattern of EMG activity in agonist and antagonist muscles: an agonist burst (AG1) initiated the development of force in the desired direction while a reciprocal burst in the antagonist (ANT-R) led to the deceleration of the force trajectory prior to the peak force. The temporal pattern of agonist and antagonist activation was dependent on force rise time. Force trajectories with long rise times (greater than 200 ms) were entirely controlled by the agonist, and EMG activity closely followed the contours of the rising force trajectory. For rise times of about 120 to 200 ms, agonist activation formed a discrete EMG burst, and force continued to rise during the subsequent silent period. For brief force rise times (less than 120 ms), reciprocal activation of the antagonist muscle occurred at about the time of the peak dF/dt. The integrated magnitude of AG1 was dependent on peak force but was independent of force rise time. AG1 duration varied directly with both peak force and force rise time. The integrated value of ANT-R varied as an inverse function of force rise time and was minimally influenced by peak force. ANT-R was present with the same magnitude and timing in both force impulses and steps when rise times were equal; therefore it did not serve to return force to baseline. Rather it served to truncate the rising force when very brief rise times were required, thus compensating for the low-pass filter properties of the agonist muscle. Subjects were able to voluntarily suppress ANT-R in rapidly accelerated force trajectories, indicating that the linkage between the commands controlling agonist and antagonist is not obligatory; however AG1 was then prolonged. Our findings emphasize that neuronal commands to opposing muscles acting at a joint must be adapted to constraints imposed by the properties of the neuromuscular plant.

Adult↗
PubMed1987

Trajectory control in targeted force impulses. II. Pulse height control.

The present report examines the control strategy adopted by subjects to modulate the amplitude of transient force responses aimed to a target. Previous studies (Freund and Budingen 1978; Ghez and Vicario 1978) suggest that subjects modulate the rate of rise of force while maintaining force rise time at a near-constant value, independent of peak force. Such studies, however, have examined only the most rapid responses where force rise time could have been at a physiological limit. We now examine whether this control policy is dependent on an instruction to produce the fastest possible trajectories, or whether it is freely selected by subjects to maximize accuracy when rise time is unconstrained. We compared responses made by six subjects, under two task conditions: 1. Fast, "make the force impulse as brief as possible"; and 2. Accurate, "be as accurate as possible without regard to rise time". Subjects were trained to produce monotonic flexion force impulses at the elbow to match the amplitudes of visually presented target shifts. Targets of three different were presented in randomized order. Responses made under the Accurate condition were less variable at each target amplitude than those under the Fast condition. Under both conditions, the initial peaks of the first and second time derivatives of force, early measures of trajectory dynamics, were strongly predictive of the peak force achieved and were correlated with the required force (target amplitude). Therefore, response trajectories must have been largely preprogrammed, and, further, the degree to which the initial peak d2F/dt2 predicts the peak force achieved represents a measure of the contribution of a preplanned motor program to trajectory formation. Subjects showed two systematic differences in trajectories between conditions. First, in all subjects force rise time was greater in the Accurate condition than in the Fast condition. Second, while in the Fast condition there was a modest dependence of force rise time on peak force, in the Accurate condition this dependence disappeared. Thus, when subjects were attempting to be as accurate as possible, they more consistently regulated force rise time around a constant value. This pulse height control policy allows responses of different amplitudes to be produced by proportional scaling of a stereotyped waveform. We conclude that a pulse height control policy with regulation of force rise time is a strategy adopted by subjects to simplify accurate control of response amplitude.

Adult↗
PubMed1987

Trajectory control in targeted force impulses. III. Compensatory adjustments for initial errors.

In the preceding study (Gordon and Ghez 1987), we showed that accurately targeted isometric force impulses produced by human subjects are governed by a pulse height control policy. Different peak forces were achieved by modulating the rate of rise of force while force rise time was maintained close to a constant value and independent of peak force. An early measure of the rate of rise of force, peak d2F/dt2, was scaled to the required force (target amplitude) and highly predictive of the peak force achieved. In six subjects examined, peak d2F/dt2 accounted for between 70% and 96% of the total variance in peak force. In the present study, we further examined these targeted responses to determine whether the residual variability not predicted by peak d2F/dt2 could be accounted for by adjustments to the force trajectories which compensated for initial errors in the scaling of the d2F/dt2. A statistical model of the determinants of peak force was tested. This model included two paths by which the target amplitude could independently influence the peak force achieved. The first path was preprogrammed pulse height control. In this path, target amplitude determined the initial rate of rise of force (peak d2F/dt2) which in turn determined the final peak force achieved. The second path was an independent influence of errors in the initial scaling of peak d2F/dt2 on peak force. Multiple regression analysis was performed on trajectory variables within the sets of responses by each subject in each condition to determine whether the second path contributed significantly to explaining the variance in peak force. In each subject and condition, there was a significant independent influence of error in d2F/dt2 on peak force, and the direction of this effect was to decrease the magnitudes of peak force errors. These compensatory adjustments accounted for between 1% and 14% of the total variance in peak force. Further multiple regression analyses revealed that inappropriate scaling of the initial phase of the trajectories was compensated for by shortening or prolonging the force rise time. These trajectory adjustments were in turn implemented by modulation of the timing and magnitude of the contractions in the agonist and antagonist muscles that produced the force trajectories. Because these compensatory adjustments were evident in the EMG pattern at latencies too short to be accounted for by peripheral feedback, we assume that they depend on internal monitoring of the unfolding neural commands. These internal feedback processes act in parallel with the programmed commands, both determining the force trajectory.

Adaptation, Physiological↗
PubMed1985

Task-related coding of stimulus and response in cat motor cortex.

In a previous study in the cat, we have reported that motor cortex neurons discharging before the initiation of an aimed forearm response (lead cells) are better timed to movement of a display (stimulus) than to the response. The present study was done to distinguish the coding of stimulus and response features in the discharge patterns of such early activity in motor cortex. Single neurons were recorded in the arm area of motor cortex in three cats performing the same pair of responses (forearm flexion and extension) but to display movements in either of the two directions by changing display polarity. The modulation of lead cell activity was contingent on the occurrence of the learned motor response and timed to the stimulus in all conditions. The majority of lead cells (88%, n = 50) fell into one of two distinct classes. In one class of neurons, force-direction (56%, n = 32), activity was contingent on a single direction of forelimb response (flexion or extension) and was thus independent of the direction of the display stimulus. The only muscles whose patterns matched the activity of this class of response-related neurons were forelimb flexors and extensors. In these neurons, the onset of modulation was timed to one or the other of the two stimuli according to the stimulus direction which elicited the appropriate response. Thus, the display-related input to these neurons varied according to the response required. In the second class of neurons, stimulus-direction (32%, n = 18), modulation was associated with a specific stimulus direction rather than the response direction. The pattern of activity of these neurons was similar to the pattern of EMG signals of shoulder and neck muscles during the different task conditions. The contraction of proximal and axial muscles corresponded to a second response elicited by the stimulus, namely attempts at head rotation towards the moving display and was independent of the conditioned forelimb response in both time of onset and direction. To test the possibility that stimulus-direction neurons participated in the control of head rotation we trained two of the animals to also produce isometric changes in neck torque in the direction of the moving display without making the forelimb response. The activity of stimulus-direction neurons was similarly modulated during performance of the neck task. By contrast, force-direction neurons examined during the neck task were either unmodulated or discharged after the neck response.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1984

Specialized subregions in the cat motor cortex: anatomical demonstration of differential projections to rostral and caudal sectors.

Ipsilateral cortico-cortical and thalamo-cortical projections to the cat motor cortex were determined from the locations of retrogradely labeled neurons following single small intracortical injections of HRP in area 4 gamma. These projections were also examined by studying the distribution of anterogradely transported axonal label following multiple injections of HRP or of tritiated amino acids in areas 1-2 of SI and in area 2pri (SII). The number of retrogradely labeled cells in areas 1-2 and in area 2pri differed markedly between HRP injection sites located in the precruciate (anterior sigmoid gyrus) and postcruciate (posterior sigmoid gyrus) subregions of area 4 gamma. These associational projections from primary and secondary somatosensory cortices were dense to postcruciate subregions but weak to the precruciate subregions. The associational projections from areas 1-2 and from area 2pri to the postcruciate subregion of area 4 gamma were topographically organized, but no clear topographic organization could be demonstrated for the precruciate projection. Anterograde terminal labeling following injection of either HRP or tritiated amino acids into areas 1-2 and area 2pri confirmed the preferential projection of somatosensory cortex to the postcruciate subregion of motor cortex. The projection from somatosensory areas 1-2 was uniform over its terminal field, but that from area 2pri was more patchy and complex. HRP injections in area 4 gamma gave rise to lamellae of labeled neurons in the ventrolateral nucleus of thalamus (VL). A topographic relationship was found between the site of injection and the location of the lamella of labeled neurons. The percentage of retrogradely labeled neurons in the shell zone surrounding the border of the ventrolateral nucleus and the ventrobasal complex (VB) was greater following postcruciate than precruciate injections, whereas fewer retrogradely labeled neurons were found in central lateral nucleus (CL) after postcruciate injections than after precruciate injections. These observations support the hypothesis that differential cortical and thalamic projections to different subregions of area 4 gamma may give rise to the different physiological properties of neurons observed in these subregions (Vicario et al. 1983; Martin et al. 1981).

Afferent Pathways↗
PubMed1984

The control of rapid limb movement in the cat. IV. Updating of ongoing isometric responses.

In cats trained to track a moving display by making rapid, isometric force adjustments, responses are characterized by extremely short reaction times (60-70 ms) and a stereotyped temporal configuration. The animal uses early derivatives of display movement to scale force responses to target stimuli of different sizes according to a learned relationship between initial display motion and required force (Ghez and Vicario 1978a, 1978b). In the present study we altered that relationship by using double stimulation and delayed feedback to assess the animals' ability to update their responses. In experiments where a second target stimulus followed the first after a controlled interval (15-120 ms) on random trials, the animal modified its response in the appropriate direction with little or no increase in reaction time. When the second stimulus called for a return to baseline, the animal aborted the ongoing response. When the second stimulus called for a doubling of force, the animal increased its phasic force output; however, this increase was not sufficient to reach the new target level and late responses were emitted. The control response which followed each experimental double stimulation trial showed consistent differences from other controls in the amplitude of both peak force and peak dF/dt. Control responses following trials calling for a return were reduced in size; those following stimuli requiring response doubling were increased. We concluded that the experimental trials not only elicited modification of ongoing responses but also caused the animal to alter its internalized gain function relating initial display derivatives to required force. In experiments where feedback was delayed after giving a first target stimulus such that the compensatory display failed to reflect the animal's initial response, the animal emitted a new updated response 70-80 ms after the first. The display trajectory which caused the cat to update its response on delayed feedback trial was identical to that of control trials with long reaction times. In this case, however, the information eliciting response updating had to be derived as a difference between the actual display trajectory and that expected by the animal, based on its experience with the tracking task. This suggests that the animal develops an internal model of display properties which is used to determine when a new response is required.

Animals↗
PubMed1984

EMG patterns in antagonist muscles during isometric contraction in man: relations to response dynamics.

We studied the EMG activity of biceps and triceps in human subjects during isometric force adjustments at the elbow. Rapid targeted force pulses exhibited stereotyped trajectories in which peak force was a linear function of the derivatives of force and the time to peak force was largely independent of its amplitude. These responses were associated with an alternating triphasic pattern of EMG bursts in agonist and antagonist muscles similar to that previously described for rapid limb movements. When the instructions demanded rapid force pulses, initial agonist bursts were of constant duration, and their magnitude was strongly related to peak force achieved. The timing of EMG bursts in antagonist pairs was closely coupled to the dynamics of the force trajectory, and the rising phase of the force was determined by both agonist and antagonist bursts. When peak force was kept constant and rise time systematically varied, the presence and magnitude of antagonist and late agonist bursts were dependent on the rate of rise of force, appearing at a threshold value and then increasing in proportion to this parameter. It is proposed that antagonist activity compensates for nonlinearity in muscle properties to enable the linear scaling of targeted forces which characterizes performance in this task.

Adult↗
PubMed1982

The control of rapid limb movement in the cat. III. Agonist - antagonist coupling.

Electromyographic (EMG) activity associated with rapid voluntary limb movements exhibits a characteristic "three burst pattern". The first burst is in the agonist muscle (AG1), the second is in the antagonist (ANT) and the third is again in the agonist (AG2). The present study was undertaken to determine whether ANT and AG2 reflect preprogrammed commands or responses to stretch consequent upon limb displacement. To answer this question EMG activity of agonist and antagonist muscles was examined in cats performing a tracking task. To dissociate centrally programmed muscular events from their intended mechanical consequences, isometric and anisometric conditions were presented in either a predictable or unpredictable sequence. A torque motor was used to control limb trajectory and to impose passive angular displacements. Whereas AG1 was present under both isometric and anisometric conditions, ANT and AG2 required limb displacement and were time locked to movement parameters. ANT occurred within 15 ms following the onset of acceleration. its magnitude varied linearly with this parameter and inversely with AG1. Passive displacements stretching the antagonist elicited responses with similar latencies and greatest magnitude for a given acceleration. AG2 was only present in underdamped movements with terminal oscillations and typically occurred when the position reached its peak and the velocity recrossed zero. Its magnitude was a function of both limb deceleration and of intended force. The data indicate that both ANT and AG2 represent responses to muscles stretch whose amplitudes are modulated by descending commands. Reciprocal mechanisms operating at a spinal level could account for the reduction of the antagonist response as a function of intended force. The increased sensitivity of late stretch responses in the agonist with higher intended forces is compatible with motoneuron facilitation by tonic descending commands. It is proposed that the stretch evoked responses function to dampen terminal oscillations which ensue from rapid displacement of the mass of the limb against elastic forces of muscle and soft tissue.

Animals↗
PubMed1978

The control of rapid limb movement in the cat. I. Response latency.

A versatile tracking task has been developed to assess the competence of feline subjects in using sensory information to initiate and guide limb movement. In the present study we investigated the response latency and the factors which affect it in order to define temporal constraints on the underlying processing of information. The paradigm makes use of an electromechanical display of the difference between the output of transducers in a manipulandum (controlled by the cat) and a target level (controlled by the experimenter). Because of inertia and friction, a step change in target level required 230 msec to fully shift the display. The animals were trained to match the position of the manipulandum (or the force they applied to it isometrically) to the target level for a food reward. Target perturbations elicited rapid position or force adjustments of appropriate direction and magnitude at short latency. Under isometric conditions, the latency of the response was an inverse function of the extent of display motion and of the peak rate of force change attained during the adjustment. Asymptotic values of the response time, measured from the perturbation to the first change in force applied to the lever, were typically between 50 and 70 msec. Since the motor responses were invariably initiated while the display was still moving, the animals must have used information contained in the derivatives of its motion. In the absence of visual cues, deflection of the vibrissae by the sudden motion of the dispaly provided two of the four animals with sufficient information to initiate and guide their movements without changes in response latency. When vision was allowed, section of the vibrissae produced a persistent increase in response times of 15-20 msec. This difference in latency is compatible with the delays due to retinal transmission. The short time interval elapsing between stimulus and response strongly suggests that the topography and the metrics of the underlying transformations are adjusted by gating and/or biasing processes which precede the stimulus. It is concluded that sensory information can generate purposeful movements through pathways which include only a small number of central relays in series, independently of its modality.

Animals↗
PubMed1978

The control of rapid limb movement in the cat. II. Scaling of isometric force adjustments.

In the previous study it was shown that cats were capable of making rapid and accurate adjustments in the force they applied to a lever in accord with information provided by a compensatory display. In the present study, isometric responses were examined in greater detail to determine 1. if a general control policy (or model) governing responses of different magnitudes could be inferred from the relations among output parameters and 2. if the earliest output measures were scaled to the preceding sensory events. The force adjustments elicited by the sudden motion of the display showed a linera relation between the peak force and the peak of its first derivative, dF/dt. Similarly, the peak d2F/dt2 was a linear function of dF/dt. By contrast, the times required to achieve the peak force and the peak dF/dt were largely independent of their magnitudes. These adjustments were produced by a burst of EMG activity in agonist muscles which coincided with the rising phase of dF/dt. The observations suggest that such motor outputs are determined by a pulse-step control policy. The amplitude of the pulse would control the rate of rise of dF/dt (and therefore also the peak force since the rising phase of dF/dt was of constant duration), and the step would control the level of the terminal steady state force. Both the peak force and the preceding peak dF/dt were highly correlated with the amplitude of the perturbation. Changes in display gain, which altered the required relation between input and output magnitudes, resulted in a gradual readjustment of the output parameters. It was concluded that the motor outputs were scaled from their inception to requirements dictated by the initial sensory information. The selection by the cat of the appropriate scaling function was contingent upon its previous experience with the device.

Animals↗
PubMed1978

Spinal mechanisms of the functional stretch reflex.

A sudden and rapid angular displacement of the limb evokes, in human and monkey subjects, a segmented pattern of electromyographic activity in muscles which are stretched. While the first segment is acknowledged to represent a tendon jerk, it has been proposed that the second segment, occurring with a shorter latency than a reaction time, is mediated by a transcortical loop. The present experiments were conducted in cats to determine the properties of muscle responses to torque perturbations analogous to those used in the monkey, and to determine if the integrity of supraspinal pathways is required for the individual response segments to occur. Torque perturbations which flexed the forearm evoked a segmented response in the electromyogram of the cat triceps muscle. This response typically consisted of three early segments with latencies of 10, 30 and 60 msec which were similar to the M1, M2, and M3 segments described in the monkey. The M3 and occasionally M2 components were depressed when the cat followed rather than resisted the perturbation. A torque pulse of 10 msec duration was sufficient to elicit a near maximal M1 response while torque pulses in excess of 20 msec were required to evoke the M2 response. To determine if any of these components required mediation by the cerebral cortex, experiments were conducted in decerebrate and spinal cats. Similar torque perturbations produced segmented electromyographic responses in the triceps muscles which were indistinguishable in their timing from those observed in intact cats. The torque required to produce the segmented responses was comparable as well. All three segments were dependent upon the activation of receptors in the homonymous muscle and did not require cutaneous input. These observations show that receptor properties and/or spinal mechanisms involved in the stretch reflex are sufficient to produce a segmented response similar to that observed in intact animals.

Animals↗
PubMed1978

Discharge of red nucleus neurons during voluntary muscle contraction: activity patterns and correlations with isometric force.

Discharges of red nucleus neurons were analyzed in the cat during voluntary muscle contractions performed under isometric and anisometric conditions. The observations established: (1) that neurons of the red nucleus modulate their firing in advance of the motor output under both conditions; (2) that increased activity of these neurons is specific to the direction of the force exerted ; (3) that this increase is primarily related to the magnitude of the rate of force change generated by the animal.

Action Potentials↗
PubMed1977

Spinal branching of rubrospinal axons in the cat.

The branching patterns of rubrospinal (RS) axons projecting to the cervical spinal cord between C3 and C8 were studied in the cat. RS neurons were identified by their antidromic responses to microstimulation of local axon branches within the cervical gray matter. Twenty-six of 58 RS neurons projecting to the cervical gray matter also sent axon branches to the thoracic spinal cord. Two out of 40 of these RS neurons also sent axon branches to the lumbar spinal cord. Using a collision technique, it was demonstrated that stem axons of rubrospinal neurons commonly sent multiple collaterals to different cervical segments. Neurons projecting to the cervical spinal cord alone were located in the dorsal quadrants of the red nucleus. Those projecting to cervical, as well as to more caudal segments, were intermingled with the former, and in slightly more ventral portions of the red nucleus. The presence of RS neurons projecting to widely separate levels of the spinal cord suggests that individual RS neurons may be capable of ultimately influencing two or more different motoneuron pools.

Animals↗
PubMed1975

Input-output relations of the red nucleus in the cat.

In unanesthetized cats, microstimulation within the red nucleus produces contraction of single muscles of the contralateral limbs and face. Separate zones may activate different muscles. Forelimb muscles were primarily activated from areas in the dorsomedial quadrants of the red nucleus whereas hindlimb muscles were predominantly activated from the ventorlateral quadrants. With stimulus currents of 10 muA there was considerable overlap in the effective zones activating different muscles. In the majority of cases the minimal threshold was under 10 muA when stimilating with a 50-msec pulse train. Current thresholds for electromypgraphic changes in the muscles varied inversely with pulse frequency and train duration. When long stimulus trains were applied to the red nucleus, the resulting muscle contraction was sustained for the duration of the stimulus. These motor effects did not depend upon the motor cortex or pyramidal tract but were mediated by a tract in contralateral dorsal quadrants of the spinal cord which was likely to be the rubrospinal tract. Units within the red nucleus typically had wide cutaneous receptive fields and responded to deep pressure and joint rotation in one or more limbs. Usually the focus driving the cell most briskly was located in one of the contralateral limbs and corresponded to the limb where muscle contraction was elicited by microstimulation with the same electrode. It is concluded that the red nucleus includes overlapping efferent neuronal colonies controlling individual muscles irrespective of their functional class. This property is shared by the motor cortex and suggests that these two structures may complement each other in the control of movement. The more diffuse activation of rubral than cortical neurons by natural stimuli suggests that rubral activity may not be as tightly linked as that of the motor cortex to specific peripheral input.

Animals↗