Visual and musculoskeletal underpinnings of anchoring in rhythmic visuo-motor tracking

Anchoring, that is, a local reduction in kinematic (i.e., spatio-temporal) variability, is commonly observed in cyclical movements, often at or around reversal points. Two kinds of underpinnings of anchoring have been identified—visual and musculoskeletal—yet their relative contributions and interrelations are largely unknown. We conducted an experiment to delineate the effects of visual and musculoskeletal factors on anchoring behavior in visuo-motor tracking. Thirteen participants (reduced to 12 in the analyses) tracked a sinusoidally moving visual target signal by making flexion–extension movements about the wrist, while both visual (i.e., gaze direction) and musculoskeletal (i.e., wrist posture) factors were manipulated in a fully crossed (3 × 3) design. Anchoring was affected by both factors in the absence of any significant interactions, implying that their contributions were independent. When gaze was directed to one of the target turning points, spatial endpoint variability at this point was reduced, but not temporal endpoint variability. With the wrist in a flexed posture, spatial and temporal endpoint variability were both smaller for the flexion endpoint than for the extension endpoint, while the converse was true for tracking with the wrist extended. Differential anchoring effects were absent for a neutral wrist posture and when gaze was fixated in between the two target turning points. Detailed analyses of the tracking trajectories in terms of velocity profiles and Hooke’s portraits showed that the tracking dynamics were affected more by wrist posture than by gaze direction. The discussion focuses on the processes underlying the observed independent effects of gaze direction and wrist posture on anchoring as well as their implications for the notion of anchoring as a generic feature of sensorimotor coordination.     

Local and global stabilization of coordination by sensory information

In studies of rhythmic coordination, where sensory information is often generated by an auditory stimulus, spatial and temporal variability are known to decrease at points in the movement cycle coincident with the stimulus, a phenomenon known as anchoring (Byblow et al. 1994). Here we hypothesize that the role of anchoring may be to globally stabilize coordination under conditions in which it would otherwise undergo a global coordinative change such as a phase transition. To test this hypothesis, anchoring was studied in a bimanual coordination paradigm in which either inphase or antiphase coordination was produced as auditory pacing stimuli (and hence movement frequency) were scaled over a wide range of frequencies. Two different anchoring conditions were used: a single-metronome condition, in which peak amplitude of right finger flexion coincided with the auditory stimulus; and a double-metronome condition, in which each finger reversal (flexion and extension) occurred simultaneously with the auditory stimuli. Anchored reversal points displayed lower spatial variation than unanchored reversal points, resulting in more symmetric phase plane trajectories in the double- than the single-metronome condition. The global coordination dynamics of the double-metronome condition was also more stable, with transitions from antiphase to inphase occurring less often and at higher movement frequencies than in the single-metronome condition. An extension of the Haken-Kelso-Bunz model of bimanual coordination is presented briefly which includes specific coupling of sensory information to movement through a process we call parametric stabilization. The parametric stabilization model provides a theoretical account of both local effects on the individual movement trajectories (anchoring) and global stabilization of observed coordination patterns, including the delay of phase transitions.     

The influence of joint position on the dynamics of perception-action coupling

Six right-handed subjects performed rhythmic flexion and extension movements of the index finger in time with an auditory metronome. On each block of trials, the wrist of the response hand was placed in a extended, neutral or flexed position. In the flex-on-the-beat condition, subjects were instructed to coordinate maximum excursion in the direction of finger flexion with each beat of the metronome. In the extend-on-the-beat condition, subjects were instructed to coordinate maximum excursion in the direction of finger extension with each beat of the metronome. The frequency of the metronome was increased from 2.00 Hz to 3.75 Hz in 8 steps (8 s epochs) of 0.25 Hz. During trials prepared in the extend-on-the-beat pattern, all subjects exhibited transitions to either a flex-on-the-beat pattern or to phase wandering as the frequency of pacing was increased. The time at which these transitions occurred was reliably influenced by the position of the wrist. Four subjects exhibited qualitative departures from the flex-on-the-beat pattern at pacing frequencies that were greater than those at which the extend-on-the-beat pattern could be maintained. The time at which these departures occurred was not influenced by the position of the wrist. These results are discussed with reference to the constraints imposed on the coordination dynamics by the intrinsic properties of the neuromuscular-skeletal system. Received: 1 October 1997 / Accepted: 20 March 1998     

Task-related power and coherence changes in neuromagnetic activity during visuomotor coordination

We analyzed a set of full-head (66 channels, CTF Inc.) magnetoencephalography (MEG) data recorded when 5 subjects performed rhythmic right index-finger flexion and extension movements on the beat (synchronization) or off the beat (syncopation) with a visual metronome at 1 Hz. Neuromagnetic activities in the alpha (8–14 Hz), beta (15–30 Hz) and gamma (30–50 Hz) ranges were shown to correlate with different aspects of the task. Specifically, we found that, compared with the control condition in which subjects only looked at the visual metronome without making any movement, all the movement conditions were accompanied by a decrease of power in the alpha range (8–14 Hz) in sensorimotor channels of both hemispheres, and an increase of coherence among a subset of these channels. The same comparison showed that power changes in the beta range differentiate task conditions by exhibiting power increases for synchronization and power decreases for syncopation. Changes in the gamma range power were found to be related to the kinematics of movement trajectories (flexion versus extension). These results suggest that three important cortical oscillations play different functional roles in a visuomotor timing task. Electronic Publication     

Keeping with the beat: movement trajectories contribute to movement timing

Previous studies of paced repetitive movements with respect to an external beat have either emphasised (a) the form of movement trajectories or (b) timing errors made with respect to the external beat. The question of what kinds of movement trajectories assist timing accuracy has not previously been addressed. In an experiment involving synchronisation or syncopation with an external auditory metronome we show that the nervous system produces trajectories that are asymmetric with respect to time and velocity in the out and return phases of the repeating movement cycle. This asymmetry is task specific and is independent of motor implementation details (finger flexion vs. extension). Additionally, we found that timed trajectories are less smooth (higher mean squared jerk) than unpaced ones. The degree of asymmetry in the flexion and extension movement times is positively correlated with timing accuracy. Negative correlations were observed between synchronisation timing error and the movement time of the ensuing return phase, suggesting that late arrival of the finger is compensated by a shorter return phase and conversely for early arrival. We suggest that movement asymmetry in repetitive timing tasks helps satisfy requirements of precision and accuracy relative to a target event.     

Control of aperture closure initiation during trunk-assisted reach-to-grasp movements

The present study investigated how the involvement and direction of trunk movement during reach-to-grasp movements affect the coordination between the transport and grasping components. Seated young adults made prehensile movements in which the involvement of the trunk was varied; the trunk was not involved, moved forward (flexion), or moved backward (extension) in the sagittal plane during the reach to the object. Each of the trunk movements was combined with an extension or flexion motion of the arm during the reach. Regarding the relationship between the trunk and arm motion for arm transport, the onset of wrist motion relative to that of the trunk was delayed to a greater extent for the trunk extension than for the trunk flexion. The variability of the time period from the peak of wrist velocity to the peak of trunk velocity was also significantly greater for trunk extension compared to trunk flexion. These findings indicate that trunk flexion was better integrated into the control of wrist transport than trunk extension. In terms of the temporal relationship between wrist transport and grip aperture, the relationship between the time of peak wrist velocity and the time of peak grip aperture did not change or become less steady across conditions. Therefore, the stability of temporal coordination between wrist transport and grip aperture was maintained despite the variation of the pattern of intersegmental coordination between the arm and the trunk during arm transport. The transport–aperture coordination was further assessed in terms of the control law according to which the initiation of aperture closure during the reach occurs when the hand crosses a hand-to-target distance threshold for grasp initiation, which is a function of peak aperture, wrist velocity and acceleration, trunk velocity and acceleration, and trunk-to-target distance at the time of aperture closure initiation. The participants increased the hand-to-target distance threshold for grasp initiation in the conditions where the trunk was involved compared to the conditions where the trunk was not involved. An increase also occurred when the trunk was extended compared to when it was flexed. The increased distance threshold implies an increase in the hand-to-target distance-related safety margin for grasping when the trunk is involved, especially when it is extended. These results suggest that the CNS significantly utilizes the parameters of trunk movement together with movement parameters related to the arm and the hand for controlling grasp initiation.     

Interval timing and trajectory in unequal amplitude movements

The Wing–Kristofferson (WK) model of movement timing emphasises the separation of central timer and motor processes. Several studies of repetitive timing have shown that increase in variability at longer intervals is attributable to timer processes; however, relatively little is known about the way motor aspects of timing are affected by task movement constraints. In the present study, we examined timing variability in finger tapping with differences in interval to assess central timer effects, and with differences in movement amplitude to assess motor implementation effects. Then, we investigated whether effects of motor timing observed at the point of response (flexion offset/tap) are also evident in extension, which would suggest that both phases are subject to timing control. Eleven participants performed bimanual simultaneous tapping, at two target intervals (400, 600 ms) with the index finger of each hand performing movements of equal (3 or 6 cm) or unequal amplitude (left hand 3, right hand 6 cm and vice versa). As expected, timer variability increased with the mean interval but showed only small, non-systematic effects with changes in movement amplitude. Motor implementation variability was greater in unequal amplitude conditions. The same pattern of motor variability was observed both at flexion and extension phases of movement. These results suggest that intervals are generated by a central timer, triggering a series of events at the motor output level including flexion and the following extension, which are explicitly represented in the timing system.     

Interjoint coordination during handwriting-like movements

The present study investigates intrinsic preferences and tendencies in coordination of the wrist and finger movements during handwriting-like tasks. Movement of the inkless pen tip in nine right-handed subjects was registered with a digitizer. One circle-drawing task and four line-drawing tasks were included in the experiment. The line-drawing task included: (1) drawing with the wrist only, (2) drawing with the fingers only, (3) an equivalent pattern consisting of the simultaneous flexion/extension of the wrist and fingers, and (4) a nonequivalent pattern in which wrist flexion was accompanied by finger extension and wrist extension was accompanied by finger flexion. Both the line and circle drawing were performed repetitively at four speed levels, ranging from slow to "as fast as possible" movements. The analysis of the line drawing revealed differential variability and temporal characteristics across the four movement patterns. While the equivalent pattern had characteristics of performance similar to those observed in the wrist-only and fingers-only pattern, the nonequivalent pattern was more variable and was executed slower when as fast as possible movement was required, compared to the other three patterns. The circle-drawing task also revealed intrinsic tendencies in coordination of the wrist and fingers. These tendencies were manifested by a spontaneous transition of the circular path of the pen tip to a tilted oval with increases in movement speed. The transition to the oval shape was accompanied by decreases in relative phase between the wrist and finger movements, whereas amplitudes of these movements were not affected by movement speed manipulations. The results suggest that subjects did not display a tendency to decrease the number of joints involved when executing the patterns that required simultaneous wrist and finger movements. Instead, there were preferences during these patterns to integrate wrist and finger movements with low relative phase. The findings are interpreted in terms of biomechanical constraints imposed on the wrist-finger linkage. This interpretation was further examined by testing two left-handed subjects. The data obtained showed symmetrical preferences in joint coordination. Collectively, the findings support a supposition that the shape of cursive letters may have been adjusted to the biomechanical structure of the hand to facilitate the motor act of handwriting.     

Neuromuscular-skeletal constraints upon the dynamics of perception-action coupling

Four right-handed subjects performed rhythmic flexion and extension movements of the index finger in time with an auditory metronome. On each block of trials the forearm of the response hand was placed in a prone, neutral or supine position. In the flex-on-the-beat condition, subjects were instructed to coordinate maximum excursion in the direction of finger flexion with each beat of the metronome. In the extend-on-the-beat condition, subjects were instructed to coordinate maximum excursion in the direction of finger extension with each beat of the metronome. The frequency of the metronome was increased from 1.75 Hz to 3.50 Hz in eight steps (8-s plateaus) of 0.25 Hz. During trials prepared in the extend-on-the-beat pattern, abrupt transitions to either a flex-on-the-beat pattern or to phase wandering often occurred, particularly at higher pacing frequencies. In marked constrast, during trials prepared in the flexon-the-beat pattern such transitions were never present. Both the frequency and the alacrity of these transitions were greater when the forearm was in a prone or neutral position than when the forearm was in a supine position. These results are discussed with reference to the constraints imposed on the coordination dynamics by the intrinsic properties of the neuromuscular-skeletal system.     

Reprogramming of muscle activation patterns at the wrist in compensation for elbow reaction torques during planar two-joint arm movements

The relationship between wrist kinematics, dynamics and the pattern of muscle activation were examined during a two-joint planar movement in which the two joints moved in opposite directions, i.e. elbow flexion/wrist extension and elbow extension/wrist flexion. Elbow movements (ranging from 10 to 70 deg) and wrist movements (ranging from 10 to 50 deg) were performed during a visual, step-tracking task in which subjects were required to attend to the initial and final angles at each joint. As the elbow amplitude increased, wrist movement duration increased and the wrist movement trajectories became quite variable. Analysis of the torques acting at the wrist joint showed that elbow movements produced reaction torques acting in the same direction as the intended wrist movement. Distinct patterns of muscle activation were observed at the wrist joint that were dependent on the relative magnitude of the elbow reaction torque in relation to the net wrist torque. When the magnitude of the elbow reaction torque was quite small, the wrist agonist was activated first. As the magnitude of the elbow reaction torque increased, activity in the wrist agonist decreased significantly. In conditions where the elbow reaction torque was much larger than the net wrist torque, the wrist muscle torque reversed direction to oppose the intended movement. This reversal of wrist muscle torque was directly associated with a change in the pattern of muscle activation where the wrist antagonist was activated prior to the wrist agonist. Our findings indicate that motion of the elbow joint is an important consideration in planning wrist movement. Specifically, the selection of muscle activation patterns at the wrist is dependent on the relative magnitude and direction of the elbow reaction torque in relation to the direction of wrist motion.     

Neural compensation for compliant loads during rhythmic movement

An experiment was performed to characterise the movement kinematics and the electromyogram (EMG) during rhythmic voluntary flexion and extension of the wrist against different compliant (elastic-viscous-inertial) loads. Three levels of each type of load, and an unloaded condition, were employed. The movements were paced at a frequency of 1 Hz by an auditory metronome, and visual feedback of wrist displacement in relation to a target amplitude of 100 degree was provided. Electromyographic recordings were obtained from flexor carpi radialis (FCR) and extensor carpi radialis brevis (ECR). The movement profiles generated in the ten experimental conditions were indistinguishable, indicating that the CNS was able to compensate completely for the imposed changes in the task dynamics. When the level of viscous load was elevated, this compensation took the form of an increase in the rate of initial rise of the flexor and the extensor EMG burst. In response to increases in inertial load, the flexor and extensor EMG bursts commenced and terminated earlier in the movement cycle, and tended to be of greater duration. When the movements were performed in opposition to an elastic load, both the onset and offset of EMG activity occurred later than in the unloaded condition. There was also a net reduction in extensor burst duration with increases in elastic load, and an increase in the rate of initial rise of the extensor burst. Less pronounced alterations in the rate of initial rise of the flexor EMG burst were also observed. In all instances, increases in the magnitude of the external load led to elevations in the overall level of muscle activation. These data reveal that the elements of the central command that are modified in response to the imposition of a compliant load are contingent, not only upon the magnitude, but also upon the character of the load.     

Rhythmic movements are larger and faster but with the same frequency on removal of visual feedback

The brain controls rhythmic movement through neural circuits combining visual information with proprioceptive information from the limbs. Although rhythmic movements are fundamental to everyday activities the specific details of the responsible control mechanisms remain elusive. We tested 39 young adults who performed flexion/extension movements of the forearm. We provided them with explicit knowledge of the amplitude and the speed of their movements, whereas frequency information was only implicitly available. In a series of 3 experiments, we demonstrate a tighter control of frequency compared with amplitude or speed. We found that in the absence of visual feedback, movements had larger amplitude and higher peak speed while maintaining the same frequency as when visual feedback was available; this was the case even when participants were aware of performing overly large and fast movements. Finally, when participants were asked to modulate continuously movement frequency, but not amplitude, we found the local coefficient of variability of movement frequency to be lower than that of amplitude. We suggest that a misperception of the generated amplitude in the absence of visual feedback, coupled with a highly accurate perception of generated frequency, leads to the performance of larger and faster movements with the same frequency when visual feedback is not available. Relatively low local coefficient of variability of frequency in a task that calls for continuous change in movement frequency suggests that we tend to operate at a constant frequency at the expense of variation in amplitude and peak speed.     

Neural gain induced by startling acoustic stimuli is additive to preparatory activation

Loud acoustic stimuli presented during movement preparation can shorten reaction time and increase response forcefulness. We examined how efferent connectivity of an agonist muscle to reticulospinal and corticospinal pathways, and the level of prepared movement force, affect reaction time and movement execution when the motor response is triggered by an intense acoustic stimulus. In Experiment 1, participants executed ballistic wrist flexion and extension movements of low and high force in response to visual stimuli. A loud acoustic stimulus (LAS; 105 dBa) was presented simultaneously with the visual imperative stimulus in probe trials. In Experiment 2, participants executed ballistic wrist flexion movements ranging from 10%–50% of maximum voluntary contraction with a LAS presented in probe trials. The shortening of response initiation was not affected by movement type (flexion or extension) or prepared movement force. Enhancement of response magnitude, however, was proportionally greater for low force movements and for the flexor muscle. Changes in peak force induced by the intense acoustic stimulus indicated that the neural activity introduced to motor program circuits by acoustic stimulation is additive to the voluntary neural activity that occurs due to movement preparation, rather than multiplicative.     

Individuated finger movements of rhesus monkeys: a means of quantifying the independence of the digits

1. Two rhesus monkeys were trained to perform flexion and extension movements of each digit of the right hand and of the wrist. Movements of all five digits and the wrist were monitored simultaneously. During each instructed movement, the instructed digit (or wrist) had the greatest excursion; other, noninstructed digits moved to varying degrees. 2. To assess the degree of independence of the different digits during these movements, I plotted, as a function of the instructed digit's position, the position of each noninstructed digit. The resulting trajectories typically were linear, with consistent slopes from trial to trial. 3. The slopes of these noninstructed digit versus instructed digit trajectories were used to calculate an individuation index for each instructed movement and a stationarity index for each digit. These indexes quantified two different aspects of independence. The individuation index reflects the degree to which other digits remained still during instructed movement of a given digit. The stationarity index reflects the degree to which a given digit remained still whenever it was a noninstructed digit. 4. In accordance with casual observation, thumb flexion and wrist flexion and extension consistently had both high individuation and stationarity and therefore can be said to be independent of the fingers. Although the same cannot be said of the other fingers, the present analysis provides a means of quantifying the degree of independence of these digits as well. 5. Factors are discussed that might contribute to the motion of noninstructed digits and to the trajectory linearity.     

Passive Stiffness of Coupled Wrist and Forearm Rotations

Coordinated movement requires that the neuromuscular system account and compensate for movement dynamics. One particularly complex aspect of movement dynamics is the interaction that occurs between degrees of freedom (DOF), which may be caused by inertia, damping, and/or stiffness. During wrist rotations, the two DOF of the wrist (flexion–extension and radial–ulnar deviation, FE and RUD) are coupled through interaction torques arising from passive joint stiffness. One important unanswered question is whether the DOF of the forearm (pronation–supination, PS) is coupled to the two DOF of the wrist. Answering this question, and understanding the dynamics of wrist and forearm rotations in general, requires knowledge of the stiffness encountered during rotations involving all three DOF (PS, FE, and RUD). Here we present the first-ever measurement of the passive stiffness encountered during simultaneous wrist and forearm rotations. Using a wrist and forearm robot, we measured coupled wrist and forearm stiffness in 10 subjects and present it as a 3-by-3 stiffness matrix. This measurement of passive wrist and forearm stiffness will enable future studies investigating the dynamics of wrist and forearm rotations, exposing the dynamics for which the neuromuscular system must plan and compensate during movements involving the wrist and forearm.     

Control of human arm movements in two dimensions: paths and joint control in avoiding simple linear obstacles

In order to examine path planning and the control of redundant degrees of freedom in the human arm, the movements of the shoulder, elbow and wrist were recorded as subjects moved a pointer to a target and avoided a simple obstacle. With respect to joint control, the results show that the extra degree of freedom provided by the wrist is incorporated into target movements in a systematic manner for both large and small obstacles; it is not used only when there is no geometrical alternative. For the wrist, two strategies are apparent, depending upon the length of the obstacle. Wrist extension predominates for shorter obstacles, while flexion or extension and flexion predominate for longer obstacles. These wrist movements shorten the effective length of the distal segments (lower arm plus hand and pointer) and thus reduce the excursion required at the proximal joints. In part, they correspond to assuming the most comfortable arm configuration at each point in the new path necessitated by the obstacle and can be described by static cost functions. However, wrist extension is also used to move the hand and pointer away from the obstacle as shoulder and elbow movements carry the wrist itself towards the obstacle. Wrist flexion is also used to move the pointer tip rapidly past the obstacle. These components, which cannot be explained by static cost functions alone, confirm for the human arm the hypothesized use of redundant degrees of freedom in obstacle avoidance. With respect to path planning, the results show that the minimum distance between pointer and obstacle remains fairly constant over a large range of obstacle lengths; this relative invariance is interpreted to support the hypothesis that workspace coordinates are important for movement planning. However, minimum distance and several other path parameters do depend significantly on the orientation and location of the movement in the workspace. This inhomogeneity implies that movement planning does not occur exclusively in workspace coordinates; it suggests an influence of joint space criteria. In frontal movements, for example, the systematic decline in the minimum distance with increasing obstacle length is interpreted as a compromise reducing the amount of extra joint movement and the discomfort of arm configurations.     

The onset of voluntary reactive movement is temporally influenced by the central oscillation in action tremor caused by multiple sclerosis

The influence of oscillatory activity in the motor system on the generation of voluntary movement has been previously studied by revealing temporal coupling between voluntary movements and associated physiological or pathological tremor. The present study aims to investigate whether there is any temporal correlation between the onset of a rapid reactive movement and the action tremor at the wrist in patients with multiple sclerosis (MS). In 13 MS patients, their reactive wrist movements and tremor were simultaneously recorded during a visually cued simple reaction time task. Significant correlation was found between the tremor-related and non-tremor-related measurements of the wrist movement. The onset of reactive movement was unevenly distributed over the tremor cycle peaking at 177 degrees in the direction opposite to the reactive movement, suggesting a temporal coupling between the reactive movements and tremor. No significant difference in reaction time was found between voluntary flexion and extension movements, and no significant differences in the mean values or the standard deviations of the reaction time between the movements in-phase and out-of-phase with tremor were detected, suggesting that entrainment of the spinal motor neurons is not influenced by tremor activity. In conclusion, in MS action tremor, the timing of the initiation of a rapid voluntary movement may be influenced by the pathological oscillator at a supra-spinal level.     

The influence of peripheral load on resetting voluntary movement by cortical stimulation: importance of the induced twitch

 We studied the effects of changes in loading torque on the effectiveness of magnetic cortical stimulation in evoking phase resetting of voluntary wrist movement. Nine normal subjects were studied (five on two occasions), while making rhythmical movements of the right wrist, at their preferred rate, against extension torque loads of 0.35 Nm, 0.26 Nm and 0.18 Nm, flexion torque loads of 0.09 Nm and 0.18 Nm and without external load. The position records of individual trials were used to measure the effectiveness of resetting (resetting index: the slope of the phase-response curve) and the ”null phase”, the phase to which the trials were being reset. The loading torque had a strong influence upon both the resetting index and the null phase, generated by a constant intensity of cortical stimulation such that the largest resetting indices were obtained for movements made against the largest extension torque load (mean resetting index 0.72). The degree of resetting and null phase were related to the mean amplitude and direction of the first poststimulus position peak, which in turn was largely determined by the twitch induced by the cortical shock. The timings of the averaged poststimulus position peaks following the first were simple multiples of the prestimulus movement period. Our results indicate that loading conditions profoundly influence the effectiveness of magnetic cortical stimulation in resetting a voluntary movement and that these effects appear to be largely explicable by the changes in the muscle twitch evoked by the stimulus with the different loads. We suggest that the magnetic shock is therefore unlikely to reset voluntary movement by an action directly upon the motor programme. We propose that the main method by which magnetic cortical stimulation resets repetitive wrist movement is indirect: normal generation of repetitive wrist flexion and extension is disrupted by the cortical shock, following which afferent information related to the twitch induced is able to reset the movement. Received: 2 September 1996 / Accepted: 3 April 1997     

Timing and visual feedback constraints on repetitive finger force production

While much is known about sequential effects in motor timing, less is understood about whether movement parameters such as force show sequential dependencies. In this study, we examined the effect of timing constraints on repetitive unimanual force production sequences. Ten healthy participants produced a series of pinch grip forces in time to a metronome and to visually specified force amplitudes. Either visual feedback of force produced or the auditory metronome removed 10 s into the experimental trial, with participants performing continued responses for the remaining 20 s. In the continuation trials, a negative lag-1 autocorrelation in the inter-response intervals (IRIs) was observed as is commonly seen in motor timing tasks. However, removal of visual feedback resulted in a systematic increase in mean force output through the course of the trial, resulting in positive lag-1 autocorrelation values. An interaction was found between mean IRI and peak force (PF) magnitude, with greater force variability seen for the larger intervals. However, the imposition of dual force and timing constraints had no effect either on the underlying variability of the PF or on the IRIs. The results are discussed in the context of force and time being independently specified components of a generalized motor program.     

Human finger independence: limitations due to passive mechanical coupling versus active neuromuscular control

We studied the extent to which mechanical coupling and neuromuscular control limit finger independence by studying passive and active individuated finger movements in healthy adults. For passive movements, subjects relaxed while each finger was rotated into flexion and extension by a custom-built device. For active movements, subjects moved each finger into flexion and extension while attempting to keep the other, noninstructed fingers still. Active movements were performed through approximately the same joint excursions and at approximately the same speeds as the passive movements. We quantified how mechanical coupling limited finger independence from the passive movements, and quantified how neuromuscular control limited finger independence using an analysis that subtracted the indices obtained in the passive condition from those obtained in the active condition. Finger independence was generally similar during passive and active movements, but showed a trend toward less independence in the middle, ring, and little fingers during active, large-arc movements. Mechanical coupling limited the independence of the index, middle, and ring fingers to the greatest degree, followed by the little finger, and placed only negligible limitations on the independence of the thumb. In contrast, neuromuscular control primarily limited the independence of the ring, and little fingers during large-arc movements, and had minimal effects on the other fingers, especially during small-arc movements. For the movement conditions tested here, mechanical coupling between the fingers appears to be a major factor limiting the complete independence of finger movement.