Effects of unilateral stroke on multi-finger synergies and their feed-forward adjustments.

We explored the changes in multi-finger synergies in patients after a single cortical stroke with mild motor impairments. We hypothesized that both synergy indices and anticipatory synergy adjustments prior to the initiation of a self-paced quick action would be diminished in the patients compared to age-matched controls. The patients with history of cortical stroke, and age-matched controls (n=12 in each group) performed one-finger and multi-finger accurate force production tasks involving both steady-state and quick force pulse production. Finger interdependence (enslaving) and multi-finger synergies stabilizing total force were quantified. The stroke patients showed lower maximal finger forces, in particular in the contralesional hand, which also showed increased enslaving indices. Multi-finger synergies during steady-state force production were, however, unchanged after stroke. In contrast, a drop in the synergy index prior to the force pulse generation was significantly delayed in the stroke patients. Our results show that mild cortical stroke leads to no significant changes in multifinger synergies, but there is impairment in feed-forward adjustments of the synergies prior to a quick action, a drop in the maximal force production, and an increase in enslaving. We conclude that studies of synergies reveal two aspects of synergic control differentially affected by cortical stroke.

Synergies in the space of control variables within the equilibrium-point hypothesis.

We use an approach rooted in the recent theory of synergies to analyze possible co-variation between two hypothetical control variables involved in finger force production based on the equilibrium-point (EP) hypothesis. These control variables are the referent coordinate (R) and apparent stiffness (C) of the finger. We tested a hypothesis that inter-trial co-variation in the {R; C} space during repeated, accurate force production trials stabilizes the fingertip force. This was expected to correspond to a relatively low amount of inter-trial variability affecting force and a high amount of variability keeping the force unchanged. We used the "inverse piano" apparatus to apply small and smooth positional perturbations to fingers during force production tasks. Across trials, R and C showed strong co-variation with the data points lying close to a hyperbolic curve. Hyperbolic regressions accounted for over 99% of the variance in the {R; C} space. Another analysis was conducted by randomizing the original {R; C} data sets and creating surrogate data sets that were then used to compute predicted force values. The surrogate sets always showed much higher force variance compared to the actual data, thus reinforcing the conclusion that finger force control was organized in the {R; C} space, as predicted by the EP hypothesis, and involved co-variation in that space stabilizing total force.

Task-specific stability of abundant systems: Structure of variance and motor equivalence.

Our main goal was to test a hypothesis that transient changes in performance of a steady-state task would result in motor equivalence. We also estimated effects of visual feedback on the amount of reorganization of motor elements. Healthy subjects performed two variations of a four-finger pressing task requiring accurate production of total pressing force (F TOT) and total moment of force (M TOT). In the Jumping-Target task, a sequence of target jumps required transient changes in either F TOT or M TOT. In the Step-Perturbation task, the index finger was lifted by 1cm for 0.5s leading to a change in both F TOT and M TOT. Visual feedback could have been frozen for one of these two variables in both tasks. Deviations in the space of finger modes (hypothetical commands to individual fingers) were quantified in directions of unchanged F TOT and M TOT (motor equivalent - ME) and in directions that changed F TOT and M TOT (non-motor equivalence - nME). Both the ME and nME components increased when the performance changed. After transient target jumps leading to the same combination of F TOT and M TOT, the changes in finger modes had a large residual ME component with only a very small nME component. Without visual feedback, an increase in the nME component was observed without consistent changes in the ME component. Results from the Step-Perturbation task were qualitatively similar. These findings suggest that both external perturbations and purposeful changes in performance trigger a reorganization of elements of an abundant system, leading to large ME change. These results are consistent with the principle of motor abundance corroborating the idea that a family of solutions is facilitated to stabilize values of important performance variables.

Neural control of movement stability: Lessons from studies of neurological patients.

The concept of synergy provides a theoretical framework for movement stability resulting from the neural organization of multiple elements (digits, muscles, etc.) that all contribute to salient performance variables. Although stability of performance is obviously important for steady-state tasks leading to high synergy indices, a feed-forward drop in synergy indices is seen in preparation to a quick action (i.e., anticipatory synergy adjustments, ASAs). We review recent studies of multi-finger and multi-muscle synergies that show decreased indices of synergies and ASAs in patients with Parkinson's disease (PD) or multisystem atrophy. In PD, the impairments in synergies and ASAs are partially reversed by dopaminergic drugs, and changes in synergy indices are present even in PD patients at earliest diagnosis. Taken together, these results point at subcortical structures that are crucial for proper control of movement stability. It is timely to introduce the concept of impaired control of stability as an objective, quantifiable, and theory-based clinical descriptor of movement disorders that can increase our understanding of the neural control of movement with all of its implications for clinical practice.

Challenging gait leads to stronger lower-limb kinematic synergies: The effects of walking within a more narrow pathway.

Previous studies using the uncontrolled manifold (UCM) analysis demonstrated that during the swing phase of gait, multi-joint kinematic synergies act to stabilize, i.e., minimize the variance of, the mediolateral trajectory of the swinging limb. Importantly, these synergies are strongest during midswing, suggesting that during gait, individuals may employ strategies to avoid collisions between the limbs at this instance. The purpose of the current study was to test this hypothesis by quantifying whether the synergy index (ΔV) during the middle period of the swing phase of treadmill walking was affected when the width of the treadmill belt was narrowed, a task expected to increase the risk of limb collisions. Eleven healthy young adults walked on a dual-belt treadmill under two conditions: (1) dual-belt - both belts of the treadmill moved at 1.2 m/s (total width: 62.5 cm) and the subject walked with one foot on each of the moving belts and (2) single-belt - one treadmill belt moved at 1.2m/s while the other belt remained stationary and the subject walked with both feet on the moving belt (total width: 30.5 cm). During both conditions, motion capture recorded the positions of 22 passive reflective markers from which UCM analysis was used to quantify ΔV in the joint configuration space. Results indicate that ΔV during the middle-third of swing phase significantly increased by 20% during single-belt walking (p<.01). We interpret this as evidence that the stronger synergies at midswing are needed to stabilize the limb trajectory in order to reduce the risk of between-limb collisions during a period when the lower limbs are nearest each other in the frontal plane.

Moving a hand-held object: Reconstruction of referent coordinate and apparent stiffness trajectories.

This study used the framework of the referent configuration hypothesis and slow changes in the external conditions during vertical oscillation of a hand-held object to infer the characteristics of hypothetical control variables. The study had two main objectives: (1) to show that hypothetical control variables, namely, referent coordinates and apparent stiffness of vertical hand position and grip force can be measured in an experiment; and (2) to establish relation(s) between these control variables that yield the classic grip-force-load-force coupling. Healthy subjects gripped a handle and performed vertical oscillations between visual targets at one of five metronome-prescribed frequencies. A HapticMaster robot was used to induce slow changes in the vertical force applied to the handle, while the size of the handle was changed slowly leading to changes in the grip aperture. The subjects were instructed not to react to possible changes in the external forces. A linear, second-order model was used to reconstruct the referent coordinate and apparent stiffness values for each phase of the vertical oscillation cycle using across-cycle regressions. The reconstructed time profiles of the referent coordinates and apparent stiffness showed consistent trends across subjects and movement frequencies. To validate the method, these values were used to predict the vertical force and the grip force applied to the handle for movement cycles that were not utilized in the reconstruction process. Analysis of the coupling between the four variables, two referent coordinates and two apparent stiffness values, revealed a single strong constraint reflecting the coupling between the grip force and vertical force. We view these data as providing experimental support for the idea of controlling natural, multi-muscle actions with shifts in a low-dimensional set of referent coordinates.

Intentional and unintentional multi-joint movements: their nature and structure of variance.

We tested predictions of a hierarchical scheme on the control of natural movements with referent body configurations. Subjects occupied an initial hand position against a bias force generated by a HapticMaster robot. A smooth force perturbation was applied to the hand consisting of an increase in the bias force, keeping it at a new level for 5s, and decreasing it back to the bias value. When the force returned to the bias value, the arm stopped at a position different from the initial one interpreted as an involuntary movement. We then asked subjects to make voluntary movements to targets corresponding to the measured end-position of the unintentional movements. No target for hand orientation was used. The joint configuration variance was compared between intentional and unintentional movements within the framework of the uncontrolled manifold hypothesis. Our central hypothesis was that both unintentional and intentional movements would be characterized by structure of joint configuration variance reflecting task-specific stability of salient performance variables, such as hand position and orientation. The analysis confirmed that most variance at the final steady states was compatible with unchanged values of both hand position and orientation following both intentional and unintentional movements. We interpret unintentional movements as consequences of back-coupling between the actual and referent configurations at the task level. The results suggested that both intentional and unintentional movements resulted from shifts of the body referent configuration produced intentionally or as a result of the hypothesized back-coupling. Inter-trial variance signature reflects similar task-specific stability properties of the system following both types of movements, intentional and unintentional.

Fitts' Law in early postural adjustments.

We tested a hypothesis that the classical relation between movement time and index of difficulty (ID) in quick pointing action (Fitts' Law) reflects processes at the level of motor planning. Healthy subjects stood on a force platform and performed quick and accurate hand movements into targets of different size located at two distances. The movements were associated with early postural adjustments that are assumed to reflect motor planning processes. The short distance did not require trunk rotation, while the long distance did. As a result, movements over the long distance were associated with substantial Coriolis forces. Movement kinematics and contact forces and moments recorded by the platform were studied. Movement time scaled with ID for both movements. However, the data could not be fitted with a single regression: Movements over the long distance had a larger intercept corresponding to movement times about 140 ms longer than movements over the shorter distance. The magnitude of postural adjustments prior to movement initiation scaled with ID for both short and long distances. Our results provide strong support for the hypothesis that Fitts' Law emerges at the level of motor planning, not at the level of corrections of ongoing movements. They show that, during natural movements, changes in movement distance may lead to changes in the relation between movement time and ID, for example when the contribution of different body segments to the movement varies and when the action of Coriolis force may require an additional correction of the movement trajectory.

Forces and moments generated by the human arm: variability and control.

This is an exploratory study of the accurate endpoint force vector production by the human arm in isometric conditions. We formulated three common-sense hypotheses and falsified them in the experiment. The subjects (n = 10) exerted static forces on the handle in eight directions in a horizontal plane for 25 s. The forces were of 4 magnitude levels (10, 20, 30 and 40 % of individual maximal voluntary contractions). The torsion moment on the handle (grasp moment) was not specified in the instruction. The two force components and the grasp moment were recorded, and the shoulder, elbow, and wrist joint torques were computed. The following main facts were observed: (a) While the grasp moment was not prescribed by the instruction, it was always produced. The moment magnitude and direction depended on the instructed force magnitude and direction. (b) The within-trial angular variability of the exerted force vector (angular precision) did not depend on the target force magnitude (a small negative correlation was observed). (c) Across the target force directions, the variability of the exerted force magnitude and directional variability exhibited opposite trends: In the directions where the variability of force magnitude was maximal, the directional variability was minimal and vice versa. (d) The time profiles of joint torques in the trials were always positively correlated, even for the force directions where flexion torque was produced at one joint and extension torque was produced at the other joint. (e) The correlations between the grasp moment and the wrist torque were negative across the tasks and positive within the individual trials. (f) In static serial kinematic chains, the pattern of the joint torques distribution could not be explained by an optimization cost function additive with respect to the torques. Plans for several future experiments have been suggested.

Arm motion coupling during locomotion-like actions: an experimental study and a dynamic model.

We studied the coordination of arm movements in standing persons who performed an out-of-phase arm-swinging task while stepping in place or while standing. The subjects were instructed to stop one of the arms in response to an auditory signal while trying to keep the rest of the movement pattern unchanged. A significant increase was observed in the amplitude of the arm that continued swinging under both the stepping and standing conditions. This increase was similar between the right and left arms. A dynamic model was developed including two coupled nonlinear van der Pol oscillators. We assumed that stopping an arm did not eliminate the coupling but introduced a new constraint. Within the model, superposition of two factors, a command to stop the ongoing movement of one arm and the coupling between the two oscillators, has been able to account for the observed effects. The model makes predictions for future experiments.

Unpredictable elbow joint perturbation during reaching results in multijoint motor equivalence.

Motor equivalence expresses the idea that movement components reorganize in the face of perturbations to preserve the value of important performance variables, such as the hand's position in reaching. A formal method is introduced to evaluate this concept quantitatively: changes in joint configuration due to unpredictable elbow perturbation lead to a smaller change in performance variables than expected given the magnitude of joint configuration change. This study investigated whether motor equivalence was present during the entire movement trajectory and how magnitude of motor equivalence was affected by constraints imposed by two different target types. Subjects pointed to spherical and cylindrical targets both with and without an elbow joint perturbation produced by a low- or high-stiffness elastic band. Subjects' view of their arm was blocked in the initial position, and the perturbation condition was randomized to avoid prediction of the perturbation or its magnitude. A modification of the uncontrolled manifold method variance analysis was used to investigate how changes in joint configuration on perturbed vs. nonperturbed trials (joint deviation vector) affected the hand's position or orientation. Evidence for motor equivalence induced by the perturbation was present from the reach onset and increased with the strength of the perturbation after 40% of the reach, becoming more prominent as the reach progressed. Hand orientation was stabilized more strongly by motor equivalent changes in joint configuration than was three-dimensional position regardless of the target condition. Results are consistent with a recent model of neural control that allows for flexible patterns of joint coordination while resisting joint configuration deviations in directions that affect salient performance variables. The observations also fit a general scheme of synergic control with referent configurations defined across different levels of the motor hierarchy.

Stabilization of the total force in multi-finger pressing tasks studied with the 'inverse piano' technique.

When one finger changes its force, other fingers of the hand can show unintended force changes in the same direction (enslaving) and in the opposite direction (error compensation). We tested a hypothesis that externally imposed changes in finger force predominantly lead to error compensation effects in other fingers thus stabilizing the total force. A novel device, the "inverse piano", was used to impose controlled displacements to one of the fingers over different magnitudes and at different rates. Subjects (n=10) pressed with four fingers at a constant force level and then one of the fingers was unexpectedly raised. The subjects were instructed not to interfere with possible changes in the finger forces. Raising a finger caused an increase in its force and a drop in the force of the other three fingers. Overall, total force showed a small increase. Larger force drops were seen in neighbors of the raised finger (proximity effect). The results showed that multi-finger force stabilizing synergies dominate during involuntary reactions to externally imposed finger force changes. Within the referent configuration hypothesis, the data suggest that the instruction "not to interfere" leads to adjustments of the referent coordinates of all the individual fingers.

Multi-finger interaction during involuntary and voluntary single finger force changes.

Two types of finger interaction are characterized by positive co-variation (enslaving) or negative co-variation (error compensation) of finger forces. Enslaving reflects mechanical and neural connections among fingers, while error compensation results from synergic control of fingers to stabilize their net output. Involuntary and voluntary force changes by a finger were used to explore these patterns. We hypothesized that synergic mechanisms will dominate during involuntary force changes, while enslaving will dominate during voluntary finger force changes. Subjects pressed with all four fingers to match a target force that was 10% of their maximum voluntary contraction (MVC). One of the fingers was unexpectedly raised 5.0 mm at a speed of 30.0 mm/s. During finger raising the subject was instructed "not to intervene voluntarily". After the finger was passively lifted and a new steady-state achieved, subjects pressed down with the lifted finger, producing a pulse of force voluntarily. The data were analyzed in terms of finger forces and finger modes (hypothetical commands to fingers reflecting their intended involvement). The target finger showed an increase in force during both phases. In the involuntary phase, the target finger force changes ranged between 10.71 ± 1.89% MVC (I-finger) and 16.60 ± 2.26% MVC (L-finger). Generally, non-target fingers displayed a force decrease with a maximum amplitude of -1.49 ± 0.43% MVC (L-finger). Thus, during the involuntary phase, error compensation was observed--non-lifted fingers showed a decrease in force (as well as in mode magnitude). During the voluntary phase, enslaving was observed--non-target fingers showed an increase in force and only minor changes in mode magnitude. The average change in force of non-target fingers ranged from 21.83 ± 4.47% MVC for R-finger (M-finger task) to 0.71 ± 1.10% MVC for L-finger (I-finger task). The average change in mode of non-target fingers was between -7.34 ± 19.27% MVC for R-finger (L-finger task) and 7.10 ± 1.38% MVC for M-finger (I-finger task). We discuss a range of factors affecting force changes, from purely mechanical effects of finger passive lifting to neural synergic adjustments of commands to individual fingers. The data fit a recently suggested scheme that merges the equilibrium-point hypothesis (control with referent configurations) with the idea of hierarchical synergic control of multi-element systems.

Learning motor synergies by persons with Down syndrome.

Persons with Down syndrome are frequently described as 'clumsy'. The recent progress in the development of quantitative approaches to motor synergies has allowed researchers to move towards an understanding of 'clumsiness' at the level of underlying control mechanisms. This progress has also offered an opportunity to quantify changes in motor synergies that accompany improvement in the performance of motor tasks. Previous studies of our group have shown, in particular, that persons both with and without Down syndrome are able to show improvements in indices of their multi-finger synergies in tasks that require accurate production of finger forces. In particular, 3 days of practice has been shown to lead to significant improvements in indices of multi-finger synergies that stabilize the time patterns of the total force produced by the fingers of a hand. Persons with Down syndrome showed a qualitative change in their synergies that failed to stabilize the total force altogether prior to practice and became able to do so after practice. In addition, the studies have also shown that variable practice is more beneficial for the improvement of motor synergies than blocked practice. I would like to draw an optimistic conclusion that persons with Down syndrome are not inherently 'clumsy', but have a vast potential for an improvement of their motor performance. The current state of the area of motor control allows researchers and practitioners to tap into these reserves, and to use quantitative indices of changes in motor synergies with practice to optimize motor performance of these individuals.

Vertical posture and head stability in patients with chronic neck pain.

OBJECTIVE: To evaluate postural performance and head stabilization of patients with chronic neck pain. DESIGN: A single-blind comparative group study. SUBJECTS: Patients with work-related chronic neck pain (n = 9), with chronic whiplash associated disorders (n = 9) and healthy subjects (n = 16). METHODS: During quiet standing in different conditions (e.g. 1 and 2 feet standing, tandem standing, and open and closed eyes) the sway areas and the ability to maintain the postures were measured. The maximal peak-to-peak displacement of the centre of pressure and the head translation were analysed during predictable and unpredictable postural perturbations. RESULTS: Patients with chronic neck pain, in particular those with whiplash-associated disorders, showed larger sway areas and reduced ability to successfully execute more challenging balance tasks. They also displayed larger sway areas and reduced head stability during perturbations. CONCLUSION: The results show that disturbances of postural control in chronic neck pain are dependent on the aetiology, and that it is possible to quantify characteristic postural disturbances in different neck pain conditions. It is suggested that the dissimilarities in postural performance are a reflection of different degrees of disturbances of the proprioceptive input to the central nervous system and/or of the central processing of such input.

Prehension synergies: effects of object geometry and prescribed torques.

We studied the coordination of forces and moments exerted by individual digits in static tasks that required balancing an external load and torque. Subjects ( n=10) stabilized a handle with an attachment that allowed for change of external torque. Thumb position and handle width systematically varied among the trials. Each subject performed 63 tasks (7 torque values x 3 thumb locations x 3 widths). Forces and moments exerted by the digit tips on the object were recorded. Although direction and magnitude of finger forces varied among subjects, each subject used a similar multidigit synergy: a single eigenvalue accounted for 95.2-98.5% of the total variance. When task parameters were varied, regular conjoint digital force changes (prehension synergies) were observed. Synergies represent preferential solutions used by the subjects to satisfy mechanical requirements of the tasks. In particular, chain effects in force adjustments to changes in the handle geometry were documented. An increased handle width induced the following effects: (a). tangential forces remained unchanged, (b). the same tangential forces produced a larger moment T (t), (c). the increased T (t) was compensated by a smaller moment of the normal forces T(n), and (d). normal finger forces were rearranged to generate a smaller moment. Torque control is a core component of prehension synergies. Observed prehension synergies are only mechanically necessitated in part. The data support a theory of hierarchical organization of prehension synergies.

EMG discharge patterns during human grip movement are task-dependent and not modulated by muscle contraction modes: a transcranial magnetic stimulation (TMS) study.

Our previous study revealed that, during tonic muscle contraction, remarkable functional differences among intrinsic and extrinsic muscles were observed during two different grip movements, i.e., precision and power grips. To verify whether this evidence is true even under the phasic muscle contraction, magnetic stimulation was delivered over the left scalp while a normal human subject performed phasic precision or power grip responses of the right-hand fingers in a simple reaction time (SRT) paradigm. Magnetic stimulation delivered during the latent period revealed different cortico-motoneuronal excitations between the two grip responses. In particular, the contributions of extensor carpi radialis (ECR) muscle were definitely different between the two grip responses, although motor-evoked potentials (MEPs) of first dorsal interosseous (FDI) prior to, and after EMG onset of movement initiation, were not different. These results were similar to previous results obtained during tonic muscle contraction. Thus, we have concluded that the task-dependent EMG discharge pattern in finger manipulation could not be modulated by muscle contraction modes.

Structure of motor variability in marginally redundant multifinger force production tasks.

The framework of the uncontrolled manifold hypothesis (UCM hypothesis) was applied to the analysis of the structure of finger force variability during oscillatory force production tasks. Subjects produced cycles of force with one, two (index and middle), or three (index, middle, and ring) fingers acting in parallel against force sensors mounted inside a small frame. The frame could be placed on the top of a table (stable conditions) or on a 4-mm-wide supporting surface (unstable conditions). Subjects were less variable when they used two fingers than when using one finger; adding the third finger did not change indices of variability of the performance. Components of finger force variance that did (VUN) or did not (VCOMP) change the value of a particular functional variable were computed for two control hypotheses: (1) at each time, the subjects tried to stabilize the total value of force (force-control); and (2), at each time, the subjects tried to stabilize the total moment produced with respect to an axis parallel to the hand/forearm (moment-control). Most subjects showed selective stabilization of moment and destabilization of force throughout most of the force cycle, in both stable and unstable conditions. The shapes of VUN and VCOMP suggested a possibility of selective compensation of timing errors across fingers within force cycles. One subject showed different relations between VUN and VCOMP, suggesting that these relations did in fact reflect particular central strategies of solving the tasks. The UCM method is applicable to force production tasks. It allows the comparison of control hypotheses in a quantitative way and unveils central strategies of control of redundant motor systems. Within this approach, redundancy (rather, abundance) is not a problem but an inherent part of a solution for natural motor tasks.

Anticipatory postural adjustments during load catching by standing subjects.

OBJECTIVES: (1) To study differences in the generation of anticipatory postural adjustments (APAs) in arm and trunk/leg muscles prior to catching a load released either by the subject him-/herself or by the experimenter. (2) To study the importance of different mechanical characteristics of the load at impact for the generation of APAs prior to load catching. METHODS: Standing subjects were asked to catch loads dropped onto the left hand from different heights either by the experimenter or by the subject's right hand. The load mass and release height were manipulated to keep either the mass or the momentum of the load at impact constant. APAs were quantified with integral electromyographic indices. RESULTS: APAs were observed in leg, trunk and arm muscles prior to load impact for both self- and experimenter-release trials. Kinetic energy showed higher correlations with the magnitude of APA than momentum, but only in experimenter-release trials. CONCLUSIONS: Subjects can generate APAs in both arm and trunk/leg muscles in the absence of an explicit voluntary action. The relative importance of kinetic energy and momentum for defining the magnitude of APAs can reflect the difference in the sources of information used to prepare for the forthcoming perturbation during self- and experimenter-released load catch.

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.