Biomedical subjects

P L Gribble

Publications and source records attributed to P L Gribble.

17 recordsLinked to original sources

PubMed2006

Are there distinct neural representations of object and limb dynamics?

In recent studies of human motor learning, subjects learned to move the arm while grasping a robotic device that applied novel patterns of forces to the hand. Here, we examined the generality of force field learning. We tested the idea that contextual cues associated with grasping a novel object promote the acquisition and use of a distinct internal model, associated with that object. Subjects learned to produce point-to-point arm movements to targets in a horizontal plane while grasping a robotic linkage that applied either a velocity-dependent counter-clockwise or clockwise force field to the hand. Following adaptation, subjects let go of the robot and were asked to generate the same movements in free space. Small but reliable after-effects were observed during the first eight movements in free space, however, these after-effects were significantly smaller than those observed for control subjects who moved the robot in a null field. No reduction in retention was observed when subjects subsequently returned to the force field after moving in free space. In contrast, controls who reached with the robot in a NF showed much poorer retention when returning to a force field. These findings are consistent with the idea that contextual cues associated with grasping a novel object may promote the acquisition of a distinct internal model of the dynamics of the object, separate from internal models used to control limb dynamics alone.

Adolescent↗

PubMed2005

Persistence of inter-joint coupling during single-joint elbow flexions after shoulder fixation.

Single-joint elbow flexions are associated with muscle activity at the shoulder that opposes interaction torques arising from rotation of the elbow. We have previously shown that this activity is linearly related to elbow muscle torque and is robust in the presence of novel dynamic loads. Here we examined this relationship in the context of shoulder joint fixation. We tested the hypothesis that after mechanically fixing the shoulder the relationship between shoulder muscle activity and elbow muscle torque will be preserved. In contrast, proposals in which energetic variables are optimized predict that shoulder muscle activity should cease. Subjects performed single-joint elbow flexions in a horizontal plane while interacting with the KINARM robotic exoskeleton. After repeated movements with the shoulder joint fixed we observed a slight and gradual decrease in the activity of pectoralis major relative to movements in which the shoulder was free to rotate. However the strength of the coupling between the shoulder and elbow did not change after shoulder fixation. This is consistent with our previous findings and suggests that the nervous system maintains this inter-joint coupling relationship even when activity at the fixed joint is no longer needed for movement accuracy.

Adult↗

PubMed2004

Inter-joint coupling strategy during adaptation to novel viscous loads in human arm movement.

When arm movements are perturbed by a load, how does the nervous system adjust control signals to reduce error? While it has been shown that the nervous system is capable of compensating for the effects of limb dynamics and external forces, the strategies used to adapt to novel loads are not well understood. We used a robotic exoskeleton [kinesiological instrument for normal and altered reaching movements (KINARM)] to apply novel loads to the arm during single-joint elbow flexions in the horizontal plane (shoulder rotation was allowed). Loads varied in magnitude with the instantaneous velocity of elbow flexion, and were applied to the shoulder in experiment 1 (interaction loads) and the elbow in experiment 2 (direct loads). Initial exposure to both interaction and direct loads resulted in perturbations at both joints, even though the load was applied to only a single joint. Subjects tended to correct for the kinematics of the elbow joint while perturbations at the shoulder persisted. Electromyograms (EMGs) and computed muscle torque showed that subjects modified muscle activity at the elbow to reduce elbow positional deviations. Shoulder muscle activity was also modified; however, these changes were always in the same direction as those at the elbow. Current models of motor control based on inverse-dynamics calculations and force-control, as well as models based on positional control, predict an uncoupling of shoulder and elbow muscle torques for adaptation to these loads. In contrast, subjects in this study adopted a simple strategy of modulating the natural coupling that exists between elbow and shoulder muscle torque during single-joint elbow movements.

Adaptation, Physiological↗

PubMed2003

Kinematics of wrist joint flexion in overarm throws made by skilled subjects.

Previous studies of multijoint arm movements have shown that the CNS holds arm kinematics constant in different situations by predictively compensating for the effects of interaction torques. We determined whether this was also the case for wrist joint flexion in natural overarm throws performed by skilled subjects in 3D, a situation where large passive torques can occur at the wrist. Specifically, we investigated whether wrist flexion amplitudes are held constant in throws of different speeds. Joint rotations were recorded at 1,000 Hz with the search-coil technique. Contrary to a previous study on constrained 2D throwing, indirect evidence was found that in fast throws passive torques associated with forearm deceleration were exploited to increase wrist flexion velocity. This increase in wrist flexion velocity was associated with constant wrist flexion amplitudes at ball release (mean 27 degrees) for throws of different speeds. Furthermore, final wrist flexion positions after ball release were similar for a particular subject irrespective of the speed of the throw. This was associated in faster throws with increased magnitudes of wrist flexor and wrist extensor EMG activity which damped passive torques associated with forearm angular deceleration. It is concluded that wrist flexion in overarm throws of different speeds is produced by central signals which precisely control net joint torque by both exploiting and damping passive torques during different parts of the throw to keep wrist joint angular position parameters constant. As such the results show that control strategies for natural 3D throwing are different from those for constrained 2D throwing.

Arm↗

PubMed2001

Dissociation between hand motion and population vectors from neural activity in motor cortex.

The population vector hypothesis was introduced almost twenty years ago to illustrate that a population vector constructed from neural activity in primary motor cortex (MI) of non-human primates could predict the direction of hand movement during reaching. Alternative explanations for this population signal have been suggested but could not be tested experimentally owing to movement complexity in the standard reaching model. We re-examined this issue by recording the activity of neurons in contralateral MI of monkeys while they made reaching movements with their right arms oriented in the horizontal plane-where the mechanics of limb motion are measurable and anisotropic. Here we found systematic biases between the population vector and the direction of hand movement. These errors were attributed to a non-uniform distribution of preferred directions of neurons and the non-uniformity covaried with peak joint power at the shoulder and elbow. These observations contradict the population vector hypothesis and show that non-human primates are capable of generating reaching movements to spatial targets even though population vectors based on MI activity do not point in the direction of hand motion.

Animals↗

PubMed2001

Compensation for the effects of head acceleration on jaw movement in speech.

Recent studies have demonstrated the ability of subjects to adjust the control of limb movements to counteract the effects of self-generated loads. The degree to which subjects change control signals to compensate for these loads is a reflection of the extent to which forces affecting movement are represented in motion planning. Here, we have used empirical and modeling studies to examine whether the nervous system compensates for loads acting on the jaw during speech production. As subjects walk, loads to the jaw vary with the direction and magnitude of head acceleration. We investigated the patterns of jaw motion resulting from these loads both in locomotion alone and when locomotion was combined with speech production. In locomotion alone, jaw movements were shown to vary systematically in direction and magnitude in relation to the acceleration of the head. In contrast, when locomotion was combined with speech, variation in jaw position during both consonant and vowel production was substantially reduced. Overall, we have demonstrated that the magnitude of load associated with head acceleration during locomotion is sufficient to produce a systematic change in the position of the jaw. The absence of variation in jaw position during locomotion with speech is thus consistent with the idea that in speech, the control of jaw motion is adjusted in a predictive manner to offset the effects of head acceleration.

Acceleration↗

PubMed2001

Relationship between cocontraction, movement kinematics and phasic muscle activity in single-joint arm movement.

Patterns of muscle coactivation provide a window into mechanisms of limb stabilization. In the present paper we have examined muscle coactivation in single-joint elbow and single-joint shoulder movements and explored its relationship to movement velocity and amplitude, as well as phasic muscle activation patterns. Movements were produced at several speeds and different amplitudes, and muscle activity and movement kinematics were recorded. Tonic levels of electromyographic (EMG) activity following movement provided a measure of muscle cocontraction. It was found that coactivation following movement increased with maximum joint velocity at each of two amplitudes. Phasic EMG activity in agonist and antagonist muscles showed a similar correlation that was observable even during the first 30 ms of muscle activation. All subjects but one showed statistically significant correlations on a trial-by-trial basis between tonic and phasic activity levels, including the phasic activity measure taken at the initiation of movement. Our findings provide direct evidence that muscle coactivation varies with movement velocity. The data also suggest that cocontraction is linked in a simple manner to phasic muscle activity. The similarity in the patterns of tonic and phasic activation suggests that the nervous system may use a simple strategy to adjust coactivation and presumably limb impedance in association with changes in movement speed. Moreover, since the pattern of tonic activity varies with the first 30 ms of phasic activity, the control of cocontraction may be established prior to movement onset.

Adult↗

PubMed2000

Compensation for loads during arm movements using equilibrium-point control.

A significant problem in motor control is how information about movement error is used to modify control signals to achieve desired performance. A potential source of movement error and one that is readily controllable experimentally relates to limb dynamics and associated movement-dependent loads. In this paper, we have used a position control model to examine changes to control signals for arm movements in the context of movement-dependent loads. In the model, based on the equilibrium-point hypothesis, equilibrium shifts are adjusted directly in proportion to the positional error between desired and actual movements. The model is used to simulate multi-joint movements in the presence of both "internal" loads due to joint interaction torques, and externally applied loads resulting from velocity-dependent force fields. In both cases it is shown that the model can achieve close correspondence to empirical data using a simple linear adaptation procedure. An important feature of the model is that it achieves compensation for loads during movement without the need for either coordinate transformations between positional error and associated corrective forces, or inverse dynamics calculations.

Algorithms↗

PubMed1999

Effects of gravitational load on jaw movements in speech.

External loads arising as a result of the orientation of body segments relative to gravity can affect the achievement of movement goals. The degree to which subjects adjust control signals to compensate for these loads is a reflection of the extent to which forces affecting motion are represented neurally. In the present study we assessed whether subjects, when speaking, compensate for loads caused by the orientation of the head relative to gravity. We used a mathematical model of the jaw to predict the effects of control signals that are not adjusted for changes to head orientation. The simulations predicted a systematic change in sagittal plane jaw orientation and horizontal position resulting from changes to the orientation of the head. We conducted an empirical study in which subjects were tested under the same conditions. With one exception, empirical results were consistent with the simulations. In both simulation and empirical studies, the jaw was rotated closer to occlusion and translated in an anterior direction when the head was in the prone orientation. When the head was in the supine orientation, the jaw was rotated away from occlusion. The findings suggest that the nervous system does not completely compensate for changes in head orientation relative to gravity. A second study was conducted to assess possible changes in acoustical patterns attributable to changes in head orientation. The frequencies of the first (F1) and second (F2) formants associated with the steady-state portion of vowels were measured. As in the kinematic study, systematic differences in the values of F1 and F2 were observed with changes in head orientation. Thus the acoustical analysis further supports the conclusion that control signals are not completely adjusted to offset forces arising because of changes in orientation.

Biomechanical Phenomena↗

PubMed1999

Compensation for interaction torques during single- and multijoint limb movement.

During multijoint limb movements such as reaching, rotational forces arise at one joint due to the motions of limb segments about other joints. We report the results of three experiments in which we assessed the extent to which control signals to muscles are adjusted to counteract these "interaction torques." Human subjects performed single- and multijoint pointing movements involving shoulder and elbow motion, and movement parameters related to the magnitude and direction of interaction torques were manipulated systematically. We examined electromyographic (EMG) activity of shoulder and elbow muscles and, specifically, the relationship between EMG activity and joint interaction torque. A first set of experiments examined single-joint movements. During both single-joint elbow (experiment 1) and shoulder (experiment 2) movements, phasic EMG activity was observed in muscles spanning the stationary joint (shoulder muscles in experiment 1 and elbow muscles in experiment 2). This muscle activity preceded movement and varied in amplitude with the magnitude of upcoming interaction torque (the load resulting from motion of the nonstationary limb segment). In a third experiment, subjects performed multijoint movements involving simultaneous motion at the shoulder and elbow. Movement amplitude and velocity at one joint were held constant, while the direction of movement about the other joint was varied. When the direction of elbow motion was varied (flexion vs. extension) and shoulder kinematics were held constant, EMG activity in shoulder muscles varied depending on the direction of elbow motion (and hence the sign of the interaction torque arising at the shoulder). Similarly, EMG activity in elbow muscles varied depending on the direction of shoulder motion for movements in which elbow kinematics were held constant. The results from all three experiments support the idea that central control signals to muscles are adjusted, in a predictive manner, to compensate for interaction torques-loads arising at one joint that depend on motion about other joints.

Adult↗

PubMed1998

Independent coactivation of shoulder and elbow muscles.

The aim of this study was to examine the possibility of independent muscle coactivation at the shoulder and elbow. Subjects performed rapid point-to-point movements in a horizontal plane from different initial limb configurations to a single target. EMG activity was measured from flexor and extensor muscles acting at the shoulder (pectoralis clavicular head and posterior deltoid) and elbow (biceps long head and triceps lateral head) and flexor and extensor muscles acting at both joints (biceps short head and triceps long head). Muscle coactivation was assessed by measuring tonic levels of electromyographic (EMG) activity after limb position stabilized following the end of the movements. It was observed that tonic EMG levels following movements to the same target varied as a function of the amplitude of shoulder and elbow motion. Moreover, for the movements tested here, the coactivation of shoulder and elbow muscles was found to be independent - tonic EMG activity of shoulder muscles increased in proportion to shoulder movement, but was unrelated to elbow motion, whereas elbow and double-joint muscle coactivation varied with the amplitude of elbow movement and were not correlated with shoulder motion. In addition, tonic EMG levels were higher for movements in which the shoulder and elbow rotated in the same direction than for those in which the joints rotated in opposite directions. In this respect, muscle coactivation may reflect a simple strategy to compensate for forces introduced by multijoint limb dynamics.

Elbow↗

PubMed1998

Recent tests of the equilibrium-point hypothesis (lambda model).

The lambda model of the equilibrium-point hypothesis (Feldman & Levin, 1995) is an approach to motor control which, like physics, is based on a logical system coordinating empirical data. The model has gone through an interesting period. On one hand, several nontrivial predictions of the model have been successfully verified in recent studies. In addition, the explanatory and predictive capacity of the model has been enhanced by its extension to multimuscle and multijoint systems. On the other hand, claims have recently appeared suggesting that the model should be abandoned. The present paper focuses on these claims and concludes that they are unfounded. Much of the experimental data that have been used to reject the model are actually consistent with it.

Electromyography↗

PubMed1998

Are complex control signals required for human arm movement?

It has been proposed that the control signals underlying voluntary human arm movement have a "complex" nonmonotonic time-varying form, and a number of empirical findings have been offered in support of this idea. In this paper, we address three such findings using a model of two-joint arm motion based on the lambda version of the equilibrium-point hypothesis. The model includes six one- and two-joint muscles, reflexes, modeled control signals, muscle properties, and limb dynamics. First, we address the claim that "complex" equilibrium trajectories are required to account for nonmonotonic joint impedance patterns observed during multijoint movement. Using constant-rate shifts in the neurally specified equilibrium of the limb and constant cocontraction commands, we obtain patterns of predicted joint stiffness during simulated multijoint movements that match the nonmonotonic patterns reported empirically. We then use the algorithm proposed by Gomi and Kawato to compute a hypothetical equilibrium trajectory from simulated stiffness, viscosity, and limb kinematics. Like that reported by Gomi and Kawato, the resulting trajectory was nonmonotonic, first leading then lagging the position of the limb. Second, we address the claim that high levels of stiffness are required to generate rapid single-joint movements when simple equilibrium shifts are used. We compare empirical measurements of stiffness during rapid single-joint movements with the predicted stiffness of movements generated using constant-rate equilibrium shifts and constant cocontraction commands. Single-joint movements are simulated at a number of speeds, and the procedure used by Bennett to estimate stiffness is followed. We show that when the magnitude of the cocontraction command is scaled in proportion to movement speed, simulated joint stiffness varies with movement speed in a manner comparable with that reported by Bennett. Third, we address the related claim that nonmonotonic equilibrium shifts are required to generate rapid single-joint movements. Using constant-rate equilibrium shifts and constant cocontraction commands, rapid single-joint movements are simulated in the presence of external torques. We use the procedure reported by Latash and Gottlieb to compute hypothetical equilibrium trajectories from simulated torque and angle measurements during movement. As in Latash and Gottlieb, a nonmonotonic function is obtained even though the control signals used in the simulations are constant-rate changes in the equilibrium position of the limb. Differences between the "simple" equilibrium trajectory proposed in the present paper and those that are derived from the procedures used by Gomi and Kawato and Latash and Gottlieb arise from their use of simplified models of force generation.

Arm↗

PubMed1997

Phasic and tonic stretch reflexes in muscles with few muscle spindles: human jaw-opener muscles.

We investigated phasic and tonic stretch reflexes in human jaw-opener muscles, which have few, if any, muscle spindles. Jaw-unloading reflexes were recorded for both opener and closer muscles. Surface electromyographic (EMG) activity was obtained from left and right digastric and superficial masseter muscles, and jaw orientation and torques were recorded. Unloading of jaw-opener muscles elicited a short-latency decrease in EMG activity (averaging 20 ms) followed by a short-duration silent period in these muscles and sometimes a short burst of activity in their antagonists. Similar behavior in response to unloading was observed for spindle-rich jaw-closer muscles, although the latency of the silent period was statistically shorter than that observed for jaw-opener muscles (averaging 13 ms). Control studies suggest that the jaw-opener reflex was not due to inputs from either cutaneous or periodontal mechanoreceptors. In the unloading response of the jaw openers, the tonic level of EMG activity observed after transition to the new jaw orientation was monotonically related to the residual torque and orientation. This is consistent with the idea that the tonic stretch reflex might mediate the change in muscle activation. In addition, the values of the static net joint torque and jaw orientation after the dynamic phase of unloading were related by a monotonic function resembling the invariant characteristic recorded in human limb joints. The torque-angle characteristics associated with different initial jaw orientations were similar in shape but spatially shifted, consistent with the idea that voluntary changes in jaw orientation might be associated with a change in a single parameter, which might be identified as the threshold of the tonic stretch reflex. It is suggested that functionally significant phasic and tonic stretch reflexes might not be mediated exclusively by muscle spindle afferents. Thus, the hypothesis that central modifications in the threshold of the tonic stretch reflex underlie the control of movement may be applied to the jaw system.

Electromyography↗

PubMed1997

An examination of the degrees of freedom of human jaw motion in speech and mastication.

The kinematics of human jaw movements were assessed in terms of the three orientation angles and three positions that characterize the motion of the jaw as a rigid body. The analysis focused on the identification of the jaw's independent movement dimensions, and was based on an examination of jaw motion paths that were plotted in various combinations of linear and angular coordinate frames. Overall, both behaviors were characterized by independent motion in four degrees of freedom. In general, when jaw movements were plotted to show orientation in the sagittal plane as a function of horizontal position, relatively straight paths were observed. In speech, the slopes and intercepts of these paths varied depending on the phonetic material. The vertical position of the jaw was observed to shift up or down so as to displace the overall form of the sagittal plane motion path of the jaw. Yaw movements were small but independent of pitch, and vertical and horizontal position. In mastication, the slope and intercept of the relationship between pitch and horizontal position were affected by the type of food and its size. However, the range of variation was less than that observed in speech. When vertical jaw position was plotted as a function of horizontal position, the basic form of the path of the jaw was maintained but could be shifted vertically. In general, larger bolus diameters were associated with lower jaw positions throughout the movement. The timing of pitch and yaw motion differed. The most common pattern involved changes in pitch angle during jaw opening followed by a phase predominated by lateral motion (yaw). Thus, in both behaviors there was evidence of independent motion in pitch, yaw, horizontal position, and vertical position. This is consistent with the idea that motions in these degrees of freedom are independently controlled.

Humans↗

PubMed1996

Coarticulation of jaw movements in speech production: is context sensitivity in speech kinematics centrally planned?

Coarticulation in speech production is a phenomenon in which the articulator movements for a given speech sound vary systematically with the surrounding sounds and their associated movements. Although these variations may seem to be planned centrally, without explicit models of the speech articulators, the kinematic patterns that are attributable to central control cannot be distinguished from those that arise because of dynamics and are not represented in the underlying control signals. We address the origins of coarticulation by comparing the results of empirical and modeling studies of jaw motion in speech. The simulated kinematics of sagittal-plane jaw rotation and horizontal jaw translation are compared with the results of empirical studies in which subjects produce speech-like sequences at a normal rate and volume. The simulations examine both "anticipatory" and "carryover" coarticulatory effects. In both cases, the results show that even when no account is taken of context at the level of central control, kinematic patterns vary in amplitude and duration as a function of the magnitude of the preceding or following movement, in the same manner as that observed empirically in coarticulation. Because at least some coarticulatory effects may arise from muscle mechanics and jaw dynamics and not from central control, these factors must be considered before drawing inferences about control in coarticulation.

Humans↗

PubMed1996

Origins of the power law relation between movement velocity and curvature: modeling the effects of muscle mechanics and limb dynamics.

1. When subjects trace patterns such as ellipses, the instantaneous velocity of movements is related to the instantaneous curvature of the trajectories according to a power law-movements tend to slow down when curvature is high and speed up when curvature is low. It has been proposed that this relationship is centrally planned. 2. The arm's muscle properties and dynamics can significantly affect kinematics. Even under isometric conditions, muscle mechanical properties can affect the development of muscle forces and torques. Without a model that accounts for these effects, it is difficult to distinguish between kinematic patterns that are attributable to central control and patterns that arise because of dynamics and muscle properties and are not represented in the underlying control signals. 3. In this paper we address the nature of the control signals that underlie movements that obey the power law. We use a numerical simulation of arm movement control based on the lambda version of the equilibrium point hypothesis. We demonstrate that simulated elliptical and circular movements, and elliptical force trajectories generated under isometric conditions, obey the power law even though there was no relation between curvature and speed in the modeled control signals. 4. We suggest that limb dynamics and muscle mechanics-specifically, the springlike properties of muscles-can contribute significantly to the emergence of the power law relationship in kinematics. Thus, without a model that accounts for these effects, care must be taken when making inferences about the nature of neural control.

Arm↗