Single-finger Tasks Research Articles

Unsupervised neural decoding for concurrent and continuous multi-finger force prediction

Reliable prediction of multi-finger forces is crucial for neural-machine interfaces. Various neural decoding methods have progressed substantially for accurate motor output predictions. However, most neural decoding methods are performed in a supervised manner, i.e., the finger forces are needed for model training, which may not be suitable in certain contexts, especially in scenarios involving individuals with an arm amputation. To address this issue, we developed an unsupervised neural decoding approach to predict multi-finger forces using spinal motoneuron firing information. We acquired high-density surface electromyogram (sEMG) signals of the finger extensor muscle when subjects performed single-finger and multi-finger tasks of isometric extensions. We first extracted motor units (MUs) from sEMG signals of the single-finger tasks. Because of inevitable finger muscle co-activation, MUs controlling the non-targeted fingers can also be recruited. To ensure an accurate finger force prediction, these MUs need to be teased out. To this end, we clustered the decomposed MUs based on inter-MU distances measured by the dynamic time warping technique, and we then labeled the MUs using the mean firing rate or the firing rate phase amplitude. We merged the clustered MUs related to the same target finger and assigned weights based on the consistency of the MUs being retained. As a result, compared with the supervised neural decoding approach and the conventional sEMG amplitude approach, our new approach can achieve a higher R2 (0.77 ± 0.036 vs. 0.71 ± 0.11 vs. 0.61 ± 0.09) and a lower root mean square error (5.16 ± 0.58 %MVC vs. 5.88 ± 1.34 %MVC vs. 7.56 ± 1.60 %MVC). Our findings can pave the way for the development of accurate and robust neural-machine interfaces, which can significantly enhance the experience during human-robotic hand interactions in diverse contexts.

Synergies at the level of motor units in single-finger and multi-finger tasks.

We explored the organization of motor units recorded in the flexor digitorum superficialis into stable groups (MU-modes) and force-stabilizing synergies in spaces of MU-modes. Young, healthy participants performed one-finger and three-finger accurate cyclical force production tasks. Two wireless sensor arrays (Trigno Galileo, Delsys, Inc.) were placed over the proximal and distal portions of the muscle for surface recording and identification of motor unit action potentials. Principal component analysis with Varimax rotation and factor extraction was used to identify MU-modes. The framework of the uncontrolled manifold hypothesis was used to analyze inter-cycle variance in the space of MU-modes and compute the index of force-stabilizing synergy. Multiple linear regression between the first MU-mode in the three-finger task and the first MU-modes in the three single-finger tasks showed no differences between the data recorded by the two electrodes suggesting that MU-modes were unlikely to be synonymous with muscle compartments. Multi-MU-mode synergies stabilizing task force were documented across all tasks. In contrast, there were no force-stabilizing synergies in the three-finger task analyzed in the space of individual finger forces. Our results confirm the synergic organization of motor units in single-finger tasks and, for the first time, expand this result to multi-finger tasks. We offer an interpretation of the findings within the theoretical scheme of control with spatial referent coordinates expanded to the analysis of individual motor units. The results confirm trade-offs between synergies at different hierarchical levels and expand this notion to intra-muscle synergies.

Finger Force Matching and Verbal Reports: Testing Predictions of the Iso-Perceptual Manifold Concept

We used force matching and verbal reports of finger force to explore a prediction of the iso-perceptual manifold concept, which assumes that stable percepts are associated with a manifold in the afferent-efferent space. Young subjects produced various force magnitudes with the index finger, middle finger, or both fingers together. Further, they reported the force level using a verbal scale and by matching the force with fingers of the contralateral hand. Force matching, but not verbal reports, showed larger variable errors for individual fingers in the two-finger task compared to the single-finger tasks. We discuss possible differences in afferent and efferent contributions to force perception at low and high forces based on the idea of motor control with referent coordinates for the effectors. The differences between the force matching and verbal reports are possibly related to neural circuitry differences between perceiving without action and perceiving-to-act.

Exploring the Concept of Iso-perceptual Manifold (IPM): A Study of Finger Force-Matching Tasks

We used force-matching tasks between the two hands to test predictions of the recently introduced scheme of perception based on the concept of iso-perceptual manifold (IPM) in the combined afferent–efferent space of neural signals. The main hypothesis was that accuracy and variability of individual finger force matching would be worse in a four-finger task compared to one-finger tasks. The subjects produced accurate force levels under visual feedback by pressing with either all four fingers or by one of the fingers of a hand (task-hand). They tried to match the total four-finger force or individual finger forces by pressing with the other hand (match-hand, no visual feedback). The match-hand consistently overshot the task-hand force during single-finger matching episodes. It showed higher inter-trial force variability during single-finger matching when the task-hand performed the four-finger task compared to trials when the task-hand performed single-finger tasks. These findings confirm our main hypothesis by showing that perception of individual finger forces can vary in multi-finger tasks within a space (IPM) corresponding to veridical perception of total force. Matching hypothetical commands to fingers, rather than finger forces, could be responsible for the consistent force overshoots. Indices of inter-trial variance affecting and unaffecting total force showed strong stabilization of total force in the task-hand but not in the match-hand in support of an earlier hypothesis on the importance of visual feedback for force stabilization. No differences were seen between the right and left hands suggesting that the dynamic dominance hypothesis may not be generalizable to perceptual phenomena.

Difference of motor overflow depending on the impaired or unimpaired hand in stroke patients

The aim of this study was to investigate the patterns of contralateral motor overflow (i.e. mirror movement) between the homologous body parts on the right and left side, in stroke patients during single-finger and multi-finger maximum force production tasks. Forty subjects, including stroke (n=20) and normal subjects (n=20), participated in this study. The stroke subjects maximally pressed force sensors with their fingers in a flexed position using a single (index, middle, ring, or little) or all fingers (all 4 fingers) using the impaired (IH) or unimpaired (UIH) hand, while the non-patient subjects used their right hands for the same tasks. The maximal voluntary forces in the ipsilateral and unintended pressing forces of each contralateral finger were recorded during the tasks. The magnitude of motor overflow to the contralateral side was calculated using the index of contralateral independence (CI). During the single finger tasks, the finger CI was significantly decreased in the UIH (91%) compared with that in the IH (99%) or normal hands (99%). Likewise, the multiple finger tasks showed that the CI was significantly lower in the UIH (84%) compared with that in the IH (96%) or normal hands (99%). However, the maximal forces were significantly lower in the IH relative to those in the UIH and normal hands. These data demonstrate that stroke patients have greater motor overflow from the UIH to the IH.

Finger force enslaving and surplus in spinal cord injury patients

This study investigated the phenomena of finger enslaving, involuntary finger actions by non-intended fingers, and force deficit, smaller maximum force by all four fingers than the sum of individual finger maximum forces in individuals with cervical spinal cord injuries (SCI). A total of 16 subjects participated in this study: 8 with a cervical spinal cord injury and 8 controls. Each of the injured subjects had one paralyzed finger. The results showed that the efforts to produce force using any individual finger induced force production in all other fingers, suggesting finger force enslaving. The maximum force during the four-finger task was greater than the sum of the individual finger forces during single-finger tasks in the SCI group, which was reflected by positive force deficit, "force surplus". One may utilize these findings for rehabilitation of paralyzed fingers caused by cervical spinal injuries.

Do synergies decrease force variability? A study of single-finger and multi-finger force production

We tested a hypothesis that force production by multi-finger groups leads to lower indices of force variability as compared to similar single-finger tasks. Three experiments were performed with quick force production, steady-state force production under visual feedback, and steady-state force production without visual feedback. In all experiments, a range of force levels was used computed as percentages of the maximal voluntary contraction force for each involved finger combination. Force standard deviation increased linearly with force magnitude across all three experiments and all finger combinations. There were modest differences between multi-finger and single-finger tasks in the indices of force variability, significant only in the tasks with steady-state force production under visual feedback. When fingers acted in groups, each finger showed significantly higher force variability as compared to its single-finger task and as compared to the multi-finger group as a whole. Fingers that were not instructed to produce force also showed close to linear relations between force standard deviation and force magnitude. For these fingers, indices of force variability were much higher as compared to those computed for the forces produced by instructed fingers. We interpret the findings within a feed-forward scheme of multi-finger control with two inputs only one of which is related to the explicit task. The total force variability reflects variability in only the task-related component, while variability of the finger forces is also due to variability of the component that is not related to the task. The findings tentatively suggest that total force variability originates at an upper level of the control hierarchy in accordance to the Weber-Fechner law rather than reflects a "neural noise" at the segmental level.

Do Synergies Improve Accuracy? A Study of Speed-Accuracy Trade-Offs during Finger Force Production

We explored possible effects of negative covariation among finger forces in multifinger accurate force production tasks on the classical Fitts's speed-accuracy trade-off. Healthy subjects performed cyclic force changes between pairs of targets "as quickly and accurately as possible." Tasks with two force amplitudes and six ratios of force amplitude to target size were performed by each of the four fingers of the right hand and four finger combinations. There was a close to linear relation between movement time and the log-transformed ratio of target amplitude to target size across all finger combinations. There was a close to linear relation between standard deviation of force amplitude and movement time. There were no differences between the performance of either of the two "radial" fingers (index and middle) and the multifinger tasks. The "ulnar" fingers (little and ring) showed higher indices of variability and longer movement times as compared with both "radial" fingers and multifinger combinations. We conclude that potential effects of the negative covariation and also of the task-sharing across a set of fingers are counterbalanced by an increase in individual finger force variability in multifinger tasks as compared with single-finger tasks. The results speak in favor of a feed-forward model of multifinger synergies. They corroborate a hypothesis that multifinger synergies are created not to improve overall accuracy, but to allow the system larger flexibility, for example to deal with unexpected perturbations and concomitant tasks.

Hand digit control in children: motor overflow in multi-finger pressing force vector space during maximum voluntary force production

The aim of this study was to investigate the contralateral motor overflow in children during single-finger and multi-finger maximum force production tasks. Forty-five right handed children, 5-11 years of age produced maximum isometric pressing force in flexion or extension with single fingers or all four fingers of their right hand. The forces produced by individual fingers of the right and left hands were recorded and analyzed in four-dimensional finger force vector space. The results showed that increases in task (right) hand finger forces were linearly associated with non-task (left) hand finger forces. The ratio of the non-task hand finger force magnitude to the corresponding task hand finger force magnitude, termed motor overflow magnitude (MOM), was greater in extension than flexion. The index finger flexion task showed the smallest MOM values. The similarity between the directions of task hand and non-task hand finger force vectors in four-dimensional finger force vector space, termed motor overflow direction (MOD), was the greatest for index and smallest for little finger tasks. MOM of a four-finger task was greater than the sum of MOMs of single-finger tasks, and this phenomenon was termed motor overflow surplus. Contrary to previous studies, no single-finger or four-finger tasks showed significant changes of MOM or MOD with the age of children. We conclude that the contralateral motor overflow in children during finger maximum force production tasks is dependent upon the task fingers and the magnitude and direction of task finger forces.

MOTOR OVERFLOW IN MULTI-FINGER FORCE SPACE DURING MAXIMUM VOLUNTARY FORCE PRODUCTION OF CHILDREN

Objective: The main purpose of this study was to investigate changes in finger interaction after stroke with strongly unilateral motor effects. Effects of age on finger interaction were also analyzed.Methods: Sixteen stroke subjects and 16 control subjects produced maximal voluntary contractions with different finger combinations by one hand and by two hands simultaneously. Individual finger forces were measured. In multi-finger tasks, force deficit (FD) was quantified as the difference between the peak finger forces in single-finger tasks and in multi-finger tasks, while enslaving (ENSL) was quantified as forces produced by fingers that were not required to produce force.Results: In stroke subjects, the peak forces produced by the fingers of the impaired hand (IH) were about 36% less than those produced by the unimpaired hand. Stroke resulted in higher ENSL and decreased FD in the IH, particularly when the index and middle fingers produced force together, while aging led to higher FD and no change in ENSL. Two-hand tasks were accompanied by an additional drop in the force of individual fingers, i.e. bilateral deficit (BD). No changes in BD were observed with age or after stroke.Conclusions: We conclude that IH function in persons after stroke is accompanied not only by a general loss of finger force but also by changes in indices of multi-finger interaction. The contrast between the significantly changed indices of one-hand multi-finger interaction and unchanged BD implies that cortical neurons mediating interhemispheric inhibition are relatively spared in unilateral stroke.Significance: The study shows that stroke leads to changes not only in finger force but also in finger interaction. The conclusion on relatively spared interhemispheric projections is potentially important for therapy of hand function in stroke survivors.

Interaction between Finger Force and Neural Command in Multi-Finger Force Production

In this study, we investigated the relationship between the finger force and the neural command in multi-finger force production tasks in order to characterize the neural enslaving effect and the force-deficit effect among fingers. Seven healthy male subjects were instructed to press one, two, three and four fingers on the finger sensors as hard as possible acting in parallel in all possible combinations. Then, the finger forces in each task were recorded and analyzed to represent the neural enslaving effect and the force-deficit effect. The results confirmed that individual finger forces were smaller in multi-finger maximal voluntary contraction tasks than in single-finger tasks. The force deficit effect increased with the number of fingers involved. A mathematical model proposed in this paper based on the experimental results could explicitly describe the two effects of finger interaction by representing the relationship between the neural commands and finger forces. The present results could be useful information to understand the basic neuro-muscular mechanism in hand biomechanics and the fundamentals of intelligent hand robots.

Control of finger force direction in the flexion-extension plane

We have examined the interaction among individual finger forces in tasks that required the production of the total force by a subset of fingers in a particular direction in the flexion-extension plane. Nine subjects produced fingertip forces in a prescribed direction with a maximum voluntary contraction (MVC) effort and held the peak force for two seconds. Six finger combinations were tested, four single-finger tasks--Index (I), Middle (M), Ring (R) and Little (L)--one two-digit task (IM), and one four-digit task (IMRL). The subjects were asked to generate the finger forces in two directions, 0 degrees (perpendicular to the surface of the transducer) and 15 degrees toward the palm. In all task conditions, there were two experimental sessions, with and without visual feedback on the task force vector. The main findings were: 1. The target direction significantly affected the constant error (CE) but not the variable error (VE) while removal of the feedback resulted in an increase in VE. 2. The direction of the forces produced by fingers that were not explicitly required to produce force (enslaved fingers) depended on the target direction. 3. In multi-finger tasks, the individual fingers produced force in directions that could differ significantly from the target direction, while the resultant force pointed in the target direction. There was a negative co-variation among the deviations of the directions of the individual finger forces from the target direction. If a finger force vector deviated from the target, another finger force vector was likely to deviate in the opposite direction. We conclude that a multi-finger synergy is involved in the control of the finger force direction.

Age effects on force produced by intrinsic and extrinsic hand muscles and finger interaction during MVC tasks.

Finger-pressing forces are produced by activation of the intrinsic hand muscles, which are finger specific, and the extrinsic muscles that connect to multiple fingers. We tested a hypothesis of greater weakening of intrinsic hand muscles with age and quantified associated indexes of finger interaction such as enslaving (force production by unintended fingers) and force deficit (loss of finger force in multifinger tasks compared with single-finger tasks). Twelve young (23-35 yr old) and 12 elderly (70-95 yr old) men and women performed single-finger and four-finger maximal pressing tasks, in which force was applied at the proximal phalanges (PP, the intrinsic muscles are major focal force generators) and at the distal phalanges (DP, the extrinsic muscles are focal force generators). The decline in the peak force with age was greater at PP (30%) than at DP (19%). Larger indexes of finger interaction were observed at PP (enslaving = 17.2 +/- 9.4%, force deficit = 36.1 +/- 11.1%) than at DP (enslaving = 14.9 +/- 8.8%, force deficit = 27.7 +/- 10.8%) across ages and genders. We conclude that intrinsic hand muscles show disproportionate weakening with age. The greater indexes of finger interaction in PP tests with greater involvement of intrinsic hand muscles suggest that the finger interactions are predominantly of a central origin across ages and genders.

Force and torque production in static multifinger prehension: biomechanics and control. II. Control.

The coordination of digits during combined force/torque production tasks was further studied using the data presented in the companion paper [Zatsiorsky et al. Biol Cybern this issue, Part I]. Optimization was performed using as criteria the cubic norms of (a) finger forces, (b) finger forces normalized with respect to the maximal forces measured in single-finger tasks, (c) finger forces normalized with respect to the maximal forces measured in a four-finger task, and (d) finger forces normalized with respect to the maximal moments that can be generated by the fingers. All four criteria failed to predict antagonist finger moments when these moments were not imposed by the task mechanics. Reconstruction of neural commands: The vector of neural commands c was reconstructed from the equation c=W(-1)F, where W is the finger interconnection weight matrix and F is the vector of finger forces. The neural commands ranged from zero (no voluntary force production) to one (maximal voluntary contraction). For fingers producing moments counteracting the external torque ('agonist' fingers), the intensity of the neural commands was well correlated with the relative finger forces normalized to the maximal forces in a four-finger task. When fingers produced moments in the direction of the external torque ('antagonist' fingers), the relative finger forces were always larger than those expected from the intensity of the corresponding neural commands. The individual finger forces were decomposed into forces due to 'direct' commands and forces induced by enslaving effects. Optimization of the neural commands resulted in the best correspondence between actual and predicted finger forces. The antagonist moments are, at least in part, due to enslaving effects: strong commands to agonist fingers also activated antagonist fingers.

Central mechanisms of finger interaction during one- and two-hand force production at distal and proximal phalanges

In this study we used changes in the relative involvement of different muscle groups during force production at the distal (DT) and proximal (PR) phalanges to test and modify a hypothesis on the central organization of multi-finger control for tasks involving non-homologous elements in the two hands. Ten subjects produced maximal force with different finger combinations. Two symmetrical (PR/PR and DT/DT) and two asymmetrical (PR/DT and DT/PR) combinations of force application sites in the two hands were used. During one-hand tasks, higher forces were produced at the PR site. In multi-finger tasks, total peak force was smaller than the sum of peak forces in single-finger tasks by the involved fingers (force deficit). Force production by some fingers of a hand was accompanied by involuntary force production by other fingers (enslaving). Force deficit and enslaving were both higher at the PR site. Two-hand tasks were accompanied by an additional drop in the force of individual fingers, i.e., bilateral deficit (BD). When symmetrical sites of force production were used in the two hands, BD was lower for symmetrical finger groups than for asymmetrical groups. During tests at asymmetrical sites, BD was higher and did not depend on symmetry of involved finger groups. We conclude that within-a-hand force deficit and enslaving are likely to be of a central, neural origin. An earlier introduced hypothesis has been expanded assuming that excitatory projections to contralateral finger representations exist only for homologous elements (sub-synergies) of a multi-finger force production synergy, while only inhibitory projections connect non-homologous elements.

Bilateral deficit and symmetry in finger force production during two-hand multifinger tasks.

A comprehensive study of patterns of finger forces during one-hand and two-hand multifinger maximal force production trials was performed with particular emphasis on differences between tasks involving symmetrical and asymmetrical finger groups (symmetrical and asymmetrical tasks). Twelve healthy right-handed subjects performed maximal voluntary force production tasks with different finger combinations. Force deficit (FD) for a finger group within a hand was defined as a drop in peak force in a multifinger task as compared to the sum of individual finger peak forces in single-finger tasks. FD showed a dependence on both the number of fingers within the hand and the number of fingers in the other hand. An additional drop in peak finger forces was seen in two-hand tests (bilateral deficit, BD). BD summed over two hands was independent of the number of fingers involved in the two-hand tasks, but dependent on the distribution of fingers between the two hands. BD for a hand was larger for tasks involving fewer fingers within the hand and more fingers in the other hand. It was higher for asymmetrical tasks than for symmetrical tasks. The difference between asymmetrical and symmetrical tasks was due to the different behavior of asymmetrically involved fingers. FD was larger for asymmetrical master (explicitly involved) fingers, while forces produced involuntarily by asymmetrical slave (explicitly non-involved) fingers were larger. These differences brought down the total moment produced by both hands in the frontal plane. FD and BD are phenomena of different origin whose effects sum up. The observations have led to further development of a previously proposed double-representation, mirror-image (DoReMi) hypothesis and refinement of the neural network underlying the two-hand finger interaction.

Bilateral multifinger deficits in symmetric key-pressing tasks.

Maximal voluntary force during simultaneous bilateral and multifinger exertion has been shown to be smaller than the sum of unilateral or single-finger exertions. The goal of this study was to study the force deficit associated with bilateral multifinger tasks. Eight normal college students performed four types of maximal isometric key-pressing tasks: (1) unilateral single-finger, (2) bilateral single-finger, (3) unilateral multifinger, and (4) bilateral multifinger. Forces produced by the index (I), middle (M), ring (R), and little (L) fingers and surface electromyography (EMG) of extrinsic finger flexors were recorded. Multifinger deficit (MFD) was defined as the percentage difference between the force (or EMG) produced by a set of fingers and the sum of the forces (or EMGs) produced by the individual fingers in their unilateral single-finger tasks. Bilateral deficit (BLD) was defined as the percentage difference between the force (or EMG) produced by a set of fingers and the sum of the forces (or EMGs) produced by the finger subsets of the left and right hands. Significant BLD and MFD in force and EMG were found for all bilateral multifinger tasks and some of the bilateral single-finger tasks. Both BLD and MFD were dependent on the number of fingers involved. BLD ranged from 3% to 22.7% for force and from 8.9% to 31.0% for EMG, including bilateral single-finger and bilateral multifinger tasks. MFDs in force during bilateral I-, IM-, IMR-, and IMRL-finger tasks were 13.2%, 37.8%, 53.2%, 52.3%, respectively; and the corresponding MFDs in EMG were 11.7%, 51.3%, 67.6%, and 71.0%, respectively. BLD and MFD in EMG were found to vary in parallel with the corresponding force deficits. It was suggested that the neural ceiling effect remains the most plausible mechanism underlying the observed deficits. The central nervous system is unable to activate maximally a large number of muscle groups at the same time during tasks involving multiple body parts. During bilateral multifinger tasks, the ceiling effect may be organized hierarchically: (1) a certain limited neural drive is shared bilaterally, leading to a BLD; (2) at each hand, a certain limited neural drive is shared by multiple fingers, leading to MFD within a hand; (3) the deficits at bilateral and unilateral multifinger levels are cumulative during bilateral multifinger tasks, leading to a higher deficit associated with the tasks.

The effect of fatigue on multifinger co-ordination in force production tasks in humans.

1. This study investigated the effects of fatigue, induced by production of maximal isometric force for 60 s with four fingers, upon indices of multifinger co-ordination. 2. Measurements of individual finger forces were performed during single- and multifinger maximal force production (maximal voluntary contraction, MVC) for two sites of force application, the middle of the distal or the middle of the proximal phalanxes. Two fatiguing exercises were used, involving force production at the distal phalanxes and at the proximal phalanxes. Fourteen subjects were tested. 3. The total force in four-finger tasks dropped by about 43 % when it was produced at the site involved in the fatiguing exercise. During force production at the other site, MVC dropped by 23 %. During single-finger MVC tests, force drop with fatigue was similar across all four fingers (about -25 % of their corresponding MVCs). 4. Force production by one finger was accompanied by involuntary force production by other fingers (enslaving). Enslaving remained unchanged by fatigue when measured during force generation at the site involved in the fatiguing exercise, but increased during force production at the other site. 5. The total MVC of four fingers acting in parallel was smaller than the sum of the MVCs of these fingers in single-finger tasks (force deficit). The force deficit increased with fatigue. Force-sharing patterns during four-finger tasks showed only minor changes under fatigue. 6. These results indicate that the effects of fatigue were not limited to changes in the force-generating capabilities of the muscles. In particular, fatigue could lead to a reorganisation at a neural level that defines commands to individual fingers.

Force sharing among fingers as a model of the redundancy problem.

The aim of this study was to test Bernstein's idea that motor synergies provide solutions to the motor redundancy problem. Forces produced by individual fingers of one hand were recorded in one-, two-, three-, and four-finger tasks. The subjects (n=10) were asked to produce maximal total force (maximal voluntary contraction, MVC) and to match a ramp total force profile using different combinations of fingers. We found that individual finger forces were smaller in multifinger MVC tasks than in single-finger tasks. The deficit increased with the number of fingers involved. A saturation effect was observed: when several effectors were involved, adding a new effector did not significantly change the total force output. The data confirmed the idea that the central neural drive arriving at the level of synergies has a certain limit, a ceiling, that cannot be exceeded. The central nervous system cannot maximally activate the muscles serving all the fingers at the same time. Secondly, during the course of ramp trials, forces produced by individual fingers were linearly related to each other. Hence, a force sharing pattern was established at the beginning of the trial and did not change during the ramp period. A hypothesis is suggested that force distribution among fingers may be organized so as to minimize unnecessary rotational moment with respect to the functional longitudinal axis of the hand. Finally, in the four-finger trials, variance of the total maximal force output in ten consecutive attempts was smaller than the sum of variances of the maximal individual finger forces. The finding suggests that the control system of the motor tasks studied involves at least two levels, a central neural drive level and a synergy level. At the synergy level, an intercompensation in individual finger force production is observed.