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Biomedical subjects

J F Soechting

Publications and source records attributed to J F Soechting.

At least 19 recordsLinked to original sources

Postural dependence of muscle actions: implications for neural control.

The neural control of reaching entails the specification of a precise pattern of muscle activation distributed across the many muscles of the arm. Musculoskeletal geometry limits the possible solutions to this problem. Insight into the nature of this constraint was obtained by quantifying the postural variation in the mechanical actions of six human shoulder muscles. Estimates of muscle mechanical actions were obtained by electrically stimulating muscles to the point of contraction and recording the resulting forces and torques with a six-degree-of-freedom force-torque transducer. In a given experiment, data were obtained for up to 29 different arm postures. The mechanical actions of each muscle varied systematically with arm posture, regardless of the frame of reference used to define these actions. The nature of this dependence suggests that a relatively simple strategy can be used by the nervous system to account for the changing mechanical actions of arm muscles.

Adult

Anticipatory and sequential motor control in piano playing.

Pianists were asked to play short excerpts from several pieces on an electronic keyboard. In each piece, there were two phrases whose first few notes were played identically with the right hand. Thereafter, the two phrases were played differently. The aim of the investigation was to ascertain whether or not hand and finger kinematics diverged prior to the depression of the last common note. Such a divergence would imply an anticipatory modification of sequential movements of the hand, akin to the phenomenon of coarticulation in speech. The lack of such a divergence would imply a strictly serial organization of movement sequences with one hand, as was found previously to be the case for typing. The time at which each key was depressed and released and the speed with which the key was depressed was recorded via a MIDI interface to a laboratory computer. The motion of the right wrist and of the fingers of the right hand was recorded optoelectronically. Piano playing can invoke anticipatory modifications of hand and finger kinematics. The time at which two patterns of movements diverged varied considerably from piece to piece. Playing an ascending scale with the requirement of a "thumb-under" maneuver could evoke an anticipatory modification as much as 500 ms in advance of the last common note. In another piece, keypresses appeared to be executed in a strict serial ordering and a third piece gave results intermediate between these two extremes. We interpret the results to suggest that a strict serial execution of a movement sequence is favored as long as this is compatible with the demands of the task.

Fingers

Flexibility and repeatability of finger movements during typing: analysis of multiple degrees of freedom.

The kinematics of the hand and fingers were studied during various keystrokes in typing. These movements were defined by 17 degrees of freedom of motion, and methods were developed to identify simplifying strategies in the execution of the task. Most of the analysis was restricted to the 11 degrees of freedom of the fingers, neglecting thumb and wrist motion. Temporal characteristics of the motion were defined by computing principal components, and it was found that only a few (two to four) principal components were needed to characterize motion of each of the degrees of freedom. Hierarchical relationships among patterns within and between different degrees of freedom were identified using cluster analysis. There was a considerable amount of consistency each time a given keystroke was executed by a subject, and this repeatability may imply a reduction in the number of degrees of freedom independently controlled by the nervous system. However, there also appears to be considerable flexibility in the coordination of the many joints of the hand when examined across different keys and across different subjects.

Biomechanical Phenomena

Evaluating an integrated musculoskeletal model of the human arm.

A simplified model of the mechanical properties of muscle and of the musculoskeletal geometry was used to predict torques at the shoulder and elbow during arm movements in the sagittal plane. Subjects made movements to 20 targets spaced on the diameter of a circle centered on the initial location of the hand. Movement kinematics and the electromyographic (EMG) activity of nine shoulder and elbow muscles were recorded. Muscle force was predicted using rectified EMG activity as an input to a Hill-type model of muscle dynamics. The model also made simplifying assumptions about muscle geometry. Muscle force was then converted to torque and the individual muscle torques were weighted to provide the best fit to the joint torque computed from the kinematic data. The overall fit of the model was reasonably good, but the goodness of fit was not uniform over all movement directions. The results suggest that the assumptions about the musculo-skeletal geometry, the model of muscle dynamics, and muscles not included in the analysis all contributed to the error.

Adult

Somatosensory cortical activity in relation to arm posture: nonuniform spatial tuning.

1. Single unitary activity in primate somatosensory cortex (SI) was recorded while monkeys maintained a range of static arm postures. Unit discharge was related to parameters defining the posture of the arm by multiple linear regression techniques. 2. Two monkeys were trained to grasp a manipulandum presented at locations distributed throughout their workspace. The discharge of single units in SI was recorded for 3 s while the monkeys maintained contact with the manipulandum and the mean discharge rate over this hold time was related to the location of the hand and to the shoulder and elbow joint angles of the arm. 3. Unitary activity of 171 neurons in the proximal arm region of areas 3, 1, and 2 was recorded during the task. Of the total, 78 neurons had activity that varied with the location of the hand in space. Neuronal discharge typically varied monotonically with the target location, reaching a maximum at the borders of the work-space. The discharge rate in most of these neurons varied with both shoulder and elbow angles. 4. Discharge rate was related to the hand's location along three axes by means of a polynomial fit. In approximately half of the neurons, activity varied significantly only for displacements along a single axis in space. However, many neurons exhibited nonlinear relations between hand location along this preferred axis and discharge rate. Discharge rate did not vary for displacements of the hand in the plane perpendicular to this preferred axis (null plane). 5. In other neurons, discharge rate varied for hand displacements in a plane, i.e., along two perpendicular axes. Displacements of the hand along the axis perpendicular to this plane (null axis) did not affect the discharge rate. In only a small minority of neurons did discharge rate vary for hand displacements along all three axes in space. 6. The distribution of the sensitivity of the neural population to hand displacements along arbitrary directions in space was not uniform. On average, hand displacement along a vertical axis led to the smallest modulation of neural discharge, and displacement of the hand along the anteroposterior direction led to the largest modulation of activity.

Analysis of Variance

Task dependence of primate arm postures.

We studied the relations between arm posture and hand location as monkeys performed tasks involving three-dimensional arm movements. Two specific questions were addressed: how reliable are these relations from trial to trial and from day to day, and are the arm postures dictated only by the location of the hand or are the postures also dependent on the task being performed? Based on the variable errors in linear regressions, we found that monkeys performed the tasks in a very stereotypic fashion from trial-to-trial: for a given monkey and task, knowing the position of the hand was sufficient to determine the posture of the arm. These relations did not change from day to day; the relation between hand location and posture was stable over the course of the experiments. In some of the tasks, the postures employed by the monkeys were strikingly similar to those observed in human psychophysical studies. The relations between hand location and arm posture exhibited a large degree of task dependence. Changing the required orientation of the hand resulted in changes in the arm posture.

Animals

Use of tactile afferent information in sequential finger movements.

We have investigated how tactile afferent information contributes to the generation of sequences of skilled finger movements by anesthetizing the right index fingers of experienced typists. Subjects were asked to type phrases in which the right index finger was used only once every seven to 12 keypresses. The time at which each key was depressed was recorded with a digital timer, and the translational and rotational motion of the fingers and wrist of the right hand were recorded optoelectronically from the location of reflective markers placed on the fingers. Midway through the experiment, a local anesthetic was injected at the base of the distal phalange of the right index finger. Following digital anesthesia, error rates increased considerably, mainly due to the diminished accuracy of movements of the anesthetized finger. The typing intervals following keypresses with the anesthetized fingertip were unaffected by the removal of tactile information. When errors occurred during control trials, the intervals immediately following the errors were greatly prolonged. However, errors produced with the anesthetized right index finger did not influence the timing of subsequent keypresses, implying that lack of tactile cues affected error recognition. The movement patterns during keypresses were similar before and after digital anesthesia for some subjects, while a less pronounced flexion-extension movement was seen in other subjects. The results suggest that tactile afferent information is not essential for initiating movement segments in a sequence. Rather, they emphasize the importance of this information for ensuring movement accuracy and for detecting errors.

Adult

On the form of the internal model for reaching.

We investigated, by using simulations, possible mechanisms responsible for the errors in the direction of arm movements exhibited by deafferented patients. Two aspects of altered feedforward control were evaluated: the inability to sense initial conditions and the degradation of an internal model. A simulation which assumed no compensation for variations in initial arm configuration failed to reproduce the characteristic pattern of errors. In contrast, a simulation that assumed random variability in the generation of joint torque resulted in a distribution of handpaths which resembled some aspects of the pattern of errors exhibited by deafferented patients.

Arm

Psychophysical approaches to motor control.

A variety of experimental approaches have recently helped identify the reference frames and coordinate systems that describe the control of eye and limb movements. These descriptions apply at the behavioral level and also, despite the distributed nature of neural processing, to the population responses of different neural structures. Studies on the process of adaptation to altered environments have also provided new insights into the controlled variables for movements: although handpaths can be adapted to extrinsic demands, the adaptation is, in some cases, in an intrinsic frame of reference.

Animals

Moving effortlessly in three dimensions: does Donders' law apply to arm movement?

Donders' law, as applied to the arm, predicts that to every location of the hand in space there corresponds a unique posture of the arm as defined by shoulder and elbow angles. This prediction was tested experimentally by asking human subjects to make pointing movements to a select number of target locations starting from a wide range of initial hand locations. The posture of the arm was measured at the start and end of every movement by means of video cameras. It was found that, in general, the posture of the arm at a given hand location does depend on the starting location of the movement and that, consequently, Donders' law is violated in this experimental condition. Kinematic and kinetic factors that could account for the variations in arm posture were investigated. It proved impossible to predict the final posture of the arm purely from kinematics, based on the initial posture of the arm. One hypothesis was successful in predicting final arm postures, namely that the final posture minimizes the amount of work that must be done to transport the arm from the starting location.

Arm

Errors in kinesthetic transformations for hand apposition.

We investigated the ability of human subjects to duplicate with the left hand the spatial location of the right hand, using only kinesthetic cues. Under this experimental condition, subjects produced large and idiosyncratic errors which varied with the spatial location of the hand. Tactile cues originating from contact of the hand with a rigid surface generally led to a decrease in the variable error. However, even without tactile cues, the variable error was small compared with other tasks in which subjects are required to estimate the spatial location of the hand on the basis of kinesthetic cues. The results suggest that subjects did not derive an estimate of the location of the right hand in extrapersonal space, but rather that they performed the task within intrapersonal space.

Cues

Spatial/temporal characteristics of a motor pattern for reaching.

1. Temporal patterns of electromyographic (EMG) activity were related to the direction of fast reaching movements. Reaches were to 20 targets in the sagittal plane of the human arm. 2. The subtraction of EMG levels recorded during very slow movements to each target allowed this study to focus on the phasic aspects of complex EMGs. 3. General features of the phasic spatial/temporal patterns differed across muscles, even across muscles at the same joint. This indicates that future models of cortical to motoneuronal processing must include nonuniform space-time transformations.

Arm

Muscle activation patterns for reaching: the representation of distance and time.

1. The timing and intensity of phasic muscle activation were related to the distance of reaching movements of the human arm. We dissociated phasic components of muscle activation from complete muscle activation waveforms by subtracting waveforms obtained during very slow movements. 2. We recorded electromyographic (EMG) activity from elbow and/or shoulder muscles as standing subjects reached forward and upward to targets at four distances. Accuracy was deemphasized and no terminal corrections were allowed. In the first part of the experiment subjects were asked to move at their preferred speed. In the second part of the experiment they were asked to move using a range of speeds. 3. In the first part of the experiment subjects moved faster to more distant targets but they also increased movement time as a nearly linear function of target distance. The slope of this function was very similar across subjects. The phasic EMG waveforms for different distances appeared to be similar in shape but of variable duration. EMG time base was quantified using a correlation technique that identified the time base scale factor that best superimposed a given trace with a template. This technique revealed that the slope of the relation between EMG time base and target distance was not the same for all muscles. 4. In the second part of the experiment, where subjects moved to each target at a range of specified speeds, time base scaling was again significantly different for different muscles. The scaling differed most dramatically between anterior deltoid and medial head of triceps. 5. EMG intensity was more strongly related to movement time than to distance. We quantified the correspondence of distance and movement time to phasic EMG intensity using a multiple regression analysis of all distances and speeds, assuming a power relation. Distance exponents were positive and movement time exponents were larger and negative. This implies that movement time is more important than distance in its relation to EMG intensity.

Adult

The learning of novel finger movement sequences.

1. Experienced typists typed phrases containing words in which one isolated letter was typed with one hand, while the remaining letters were typed with the contralateral hand. 2. The translational and rotational motion of the fingers and wrist of the right hand were obtained optoelectronically from the location of reflective markers placed on the fingers. 3. Midway through the experiment, the key corresponding to the isolated letter was physically switched with another key on the keyboard, and subjects typed the letter in its new location (for 140 trials). The letter "n," typed with the right index finger, was either switched with letters normally typed with the same finger (u), with a different finger but same hand (o), with the same finger of the left hand (v), or with a different finger of the left hand (w). 4. When the words were typed normally, the interkey intervals were relatively short, and the onset of movement of the right hand began before the preceding keypress with the left hand. Thus the movement of the two hands overlapped. Furthermore, the movement to the isolated key was highly stereotypical, with little trial-to-trial variability. 5. After the transposition of keys, there were prolongations in the interkey intervals, with the largest delay occurring directly before the typing of the transposed key. Switches between homologous fingers (involving mirror movements) delayed the onset of keypresses to a lesser extent than did other switches. With practice, these delays were reduced but never reached the control level. 6. After the keyswitch, the onset of movement to the isolated key did not occur on average until after the last keypress with the contralateral hand, except when the switch involved the use of homologous fingers. In the latter case, overlapping movement of the two hands was maintained. Thus the learning of a series of discrete movements does not necessarily require that each movement segment be performed sequentially. 7. After the transposition of keys, the movement pattern and time course to a given key were similar to the movement patterns for that key observed during control trials in all conditions. Thus the learning of a series of movements may involve the use of previously learned movements under new conditions. 8. The results suggest that typing movements may be organized at several levels, including the individual keystroke and word level.

Adult

Non-uniform temporal scaling of hand and finger kinematics during typing.

We examined the manner in which the keystroke kinematics of the hand and the fingers varied with the mean rate of typing by trained typists. We used words and phrases in which only one letter was typed with the right hand and all of the remaining letters were typed using the left hand. We varied the typing rate over a threefold range (intervals between keypresses ranging from 150 ms to 500 ms) with the aid of a metronome. The results from four subjects, and three letters (n, u, and o) were analyzed. We did not find a simple scaling that could account for variations in the velocity profiles with typing rate. For some subjects and some letters, the velocities were independent of typing rate. In other instances, the kinematics did depend on typing rate, but to a much greater extent prior to the time of keypress than afterward. Sometimes the velocity profiles of all of the fingers and of the hand changed in a similar manner as the interval between keypresses was varied. In other instances only the focal movement of the hand and the finger used to press the key depended on the interval, whereas the motions of the other fingers did not. We suggest that the consistencies in the velocity profiles which we observed may simplify the problem of arranging a temporally ordered sequence of goal-directed movements.

Fingers

Parallel, interdependent channels for location and orientation in sensorimotor transformations for reaching and grasping.

1. Subjects were presented with a cylinder, whose orientation with respect to the vertical and location in space varied from trial to trial. They grasped a similar cylinder in their hands and were instructed to align the grasped cylinder with the target cylinder. In some experiments the task was performed from memory, and subjects attempted to reproduce both location and orientation of the cylinder. In others, they attempted to reproduce only its orientation, either from memory or while the cylinder was in view. 2. Multivariate linear regression analysis was used to determine persistent and variable errors in performance. This analysis related the subjects' performance (reproduced orientation) to target parameters (location and orientation). 3. We have interpreted the experimental results starting from the assumption that there are two parallel neural processes underlying reaching and grasping: one relating proximal arm motion to target location and the other relating distal hand motion to target orientation. 4. Variable errors did not vary with task conditions, even when subjects were asked to dissociate target orientation from target location by matching target orientation with the arm at the side, irrespective of the location of the target. This finding suggests that the neural transformations involving target location and target orientation are performed in parallel. 5. Persistent errors did vary with task condition. The subjects made the largest errors in matching target orientation when the target was in view, but they were asked to match its orientation at a location that differed from that of the target. These errors depended mostly on the elevation of the target and on its slant (inclination relative to the vertical). They were related to the posture of the arm in a manner that suggested that matching orientation is influenced by both extrinsic (spatial) and intrinsic (arm posture) parameters. 6. The fact that persistent errors depend on target location and on arm posture as well as on target orientation implies that the neural transformation from target orientation to hand orientation is not independent of the transformation dealing with target location.

Attention

Synergistic finger movements in a skilled motor task.

When skilled typists strike one key, typically all of the fingers of one hand are in motion simultaneously. We studied the extent to which the motion of the individual fingers was similar. Subjects were asked to type lists of words, each word designed so that only one key would be pressed by a finger of one hand, the remaining letters being typed with the other hand. Changes in the lengths (flexion-extension) and orientation (abduction-adduction) of each finger were measured and the similarity of the motion of pairs of fingers was assessed by computing correlation coefficients. For each pair of fingers, the correlation coefficients were broadly distributed, but in most instances the mean was significantly greater than zero. Adjacent fingers showed a higher degree of correlation than did non-adjacent fingers. When one of the fingers was actually used to press the key, the degree of correlation decreased substantially. The results demonstrate that in this skilled motor task, the fingers usually tend to be moved together, but they can be moved independently when the task so requires.

Fingers