When a person produces isometric force with one, two, or three fingers, the other fingers of the hand also produce a certain force. Enslaving is the involuntary force production by fingers not explicitly involved in a force-production task. This study explored the enslaving effects (EE) in multi-finger tasks in which the contributions of the flexor digitorum profundus (FDP), flexor digitorum superficialis (FDS), and intrinsic muscles (INT) were manipulated. A new experimental technique was developed that allows the redistribution of the muscle activity between the FDP, FDS, and INT muscles. In the experiment, ten subjects were instructed to perform maximal voluntary contractions with all possible one-, two-, three-, and four-finger combinations. The point of force application was changed in parallel for the index, middle, ring, and little fingers from the middle of the distal phalanx, to the distal interphalangeal joint, and then to the proximal interphalangeal joint. It was found that: (1) the EE of similar amplitude were present in various experimental conditions that involved different muscle groups for force production; (2) the EE were large on average--the slave fingers could produce forces reaching 67.5% of the maximal forces produced by themselves in a single-finger task; (3) the EE were larger for neighboring fingers; and (4) the EE were non-additive--in most cases, the EE from two or three fingers were smaller than the EE from at least one finger. EE among different muscles suggest a widespread neural interaction among the structures controlling flexor muscles in the hand as the main mechanism of finger enslaving.

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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.

During maximal voluntary contraction (MVC) with several fingers, the following three phenomena are observed: (1) the total force produced by all the involved fingers is shared among the fingers in a specific manner (sharing); (2) the force produced by a given finger in a multi-finger task is smaller than the force generated by this finger in a single-finger task (force deficit); (3) the fingers that are not required to produce any force by instruction are involuntary activated (enslaving). We studied involuntary force production by individual fingers (enslaving effects, EE) during tasks when (an)other finger(s) of the hand generated maximal voluntary pressing force in isometric conditions. The subjects (n = 10) were instructed to press as hard as possible on the force sensors with one, two, three and four fingers acting in parallel in all possible combinations. The EE were (A) large, the slave fingers always producing a force ranging from 10.9% to 54.7% of the maximal force produced by the finger in the single-finger task; (B) nearly symmetrical; (C) larger for the neighboring fingers; and (D) non-additive. In most cases, the EE from two or three fingers were smaller than the EE from at least one finger (this phenomenon was coined occlusion). The occlusion cannot be explained only by anatomical musculo-tendinous connections. Therefore, neural factors contribute substantially to the EE. A neural network model that accounts for all the three effects has been developed. The model consists of three layers: the input layer that models a central neural drive; the hidden layer modeling transformation of the central drive into an input signal to the muscles serving several fingers simultaneously (multi-digit muscles); and the output layer representing finger force output. The output of the hidden layer is set inversely proportional to the number of fingers involved. In addition, direct connections between the input and output layers represent signals to the hand muscles serving individual fingers (uni-digit muscles). The network was validated using three different training sets. Single digit muscles contributed from 25% to 50% of the total finger force. The master matrix and the enslaving matrix were computed; they characterize the ability of a given finger to enslave other fingers and its ability to be enslaved. Overall, the neural network modeling suggests that no direct correspondence exists between neural command to an individual finger and finger force. To produce a desired finger force, a command sent to an intended finger should be scaled in accordance with the commands sent to the other fingers.

Movements by a standing person are commonly associated with adjustments in the activity of postural muscles to cause a desired shift of the center of pressure (COP) and keep balance. We hypothesize that such COP shifts are controlled (stabilized) using a small set of central variables (muscle modes, M-modes), while each M-mode induces changes in the activity of a subgroup of postural muscles. The main purpose of this study has been to explore the possibility of identification of muscle synergies in a postural task using the framework of the uncontrolled manifold (UCM) hypothesis employing the following three steps in data analysis: (i) Identification of M-modes: Subjects were asked to release a load from extended arms through a pulley system, resulting in a COP shift forward prior to load release. Electromyographic (EMG) activity of eleven postural muscles on one side of the body was integrated over a 100 ms interval corresponding to the early stage of the COP shift, and subjected to a principal component (PC) analysis across multiple repetitions of each task. Three PCs were identified and associated with a 'push-back M-mode', a 'push-forward M-mode' and a 'mixed M-mode'. (ii) Calculation of the Jacobian of the system, which relates changes in the magnitude of M-modes to COP shifts using regression techniques: Subjects performed three different tasks (releasing different loads at the back, voluntarily shifting body weight forward and backward, at different speeds) to verify if the relationship between magnitudes of M-modes and COP shifts is task or direction specific. (iii) UCM analysis: Three tasks were chosen (load release in the front, arm movement forward and backward) which were associated with an early shift in COP. A manifold was identified in the M-mode space corresponding to a certain average (across trials) shift of the COP and variance per degree of freedom within the UCM (V(UCM)) and orthogonal (V(ORT)) to the UCM was computed. Across subjects, V(UCM) was significantly higher than V(ORT) when analysis at the third step was performed using a Jacobian computed based on a set of tasks associated with a COP shift in the same direction but not in the opposite direction. This result confirms our hypothesis that the M-modes work together as a synergy to stabilize a desired shift of the COP. Forward and backward COP shifts are associated with different synergies based on the same three M-modes.

A method of decomposing stabilograms into two components, termed rambling and trembling, was developed. The rambling component reveals the motion of a moving reference point with respect to which the body's equilibrium is instantantly maintained. The trembling component reflects body oscillation around the reference point trajectory. The concepts of instant equilibrium point (IEP) and discrete IEP trajectory are introduced. The rambling trajectory was computed by interpolating the discrete IEP trajectory with cubic spline functions. The trembling trajectory is found as a difference between the approximated rambling trajectory and the COP trajectory. Instant values of the trembling trajectory are negatively correlated with the values of the horizontal ground reaction force at a zero time lag. It suggests that trembling is strongly influenced by a restoring force proportional to the magnitude of COP deviation from the rambling trajectory and acts without a time delay. An increment in relative COP position per unit of the restoring force, in mm/N, was on average 1.4 ± 0.4. The contribution of rambling and trembling components in the stabilogram was ascertained. The rambling variability is approximately three times larger than the trembling variability.

The objective of the study is to examine the effects of age and gender on finger coordination. Twelve young (24 +/- 8 yr; 6 men and 6 women) and 12 elderly (75 +/- 5 yr; 6 men and 6 women) subjects performed single-finger maximal contraction [maximal voluntary contraction (MVC)], four-finger MVC, and four-finger ramp force production tasks by pressing on individual force transducers. A drop in the force of individual fingers during four-finger MVC tasks compared with single-finger MVC tasks (force deficit) was larger, whereas unintended force production by other fingers during single-finger MVC tasks (enslaving) was smaller, in elderly than in young subjects and in women than in men. Force deficit was smaller and enslaving was larger in subjects with higher peak force. During the ramp task, the difference between the variance of total force and the sum of variances of individual forces showed a logarithmic relation to the level of total force, across all subject groups. These findings suggest that indexes of finger coordination scale with force-generating capabilities across gender and age groups.

When a standing person performs a movement such that the center of gravity shifts, the activity of postural muscles adjusts to keep the balance. We assume that such adjustments are controlled using a small set of central variables, while each variable induces changes in the activity of a subgroup of postural muscles. The purpose of this study has been to identify such muscle groups (muscle modes or M-modes) and compare them across tasks and subjects. Four tasks required the subjects to release a load from extended arms leading to a center of pressure (COP) shift prior to the load release. The fifth task required an explicit COP shift by voluntary sway. Electromyographic activity of 11 postural muscles on one side of the body was integrated over a 100-ms interval corresponding to the early stage of the COP shift, and this integrated EMG activity was subjected to a principal component (PC) analysis across multiple repetitions of each task. Three PCs were identified and associated with a "push-back M-mode," a "push-forward M-mode," and a "mixed M-mode." Cluster analysis of the PC vectors across tasks and across subjects confirmed the existence of distinctive push-forward and push-back muscle groups. PC vectors were also compared across tasks and across subjects using cosines as a measure of colinearity between pairs of vectors. In general, M-modes were similar across both tasks and subjects. We conclude that shifts of the COP, whether implicit or explicit, are controlled using a small set of central variables associated with changes in the activity of robust subsets of postural muscles. These results can be used for future analysis of muscle synergies associated with postural tasks.