Multi-finger Synergies Research Articles

Two classes of action-stabilizing synergies reflecting spinal and supraspinal circuitry.

We explored force-stabilizing synergies during accurate four-finger constant force production tasks in spaces of finger modes (commands to fingers computed to account for the finger interdependence) and of motor unit (MU) firing frequencies. The main specific hypothesis was that the multifinger synergies would disappear during unintentional force drifts without visual feedback on the force magnitude, whereas MU-based synergies would be robust to such drifts. Healthy participants performed four-finger accurate cyclical force production trials followed by trials of constant force production. Individual MUs were identified in the flexor digitorum superficialis (FDS) and extensor digitorum communis (EDC). Principal component analysis was applied to motor unit frequencies to identify robust MU groups (MU-modes) with parallel scaling of the firing frequencies in FDS, in EDC, and the combined MUs of FDS + EDC. The framework of the uncontrolled manifold hypothesis was used to quantify force-stabilizing synergies when visual feedback on the force magnitude was available and 15 s after turning the visual feedback off. Removing visual feedback led to a force drift toward lower magnitudes, accompanied by the disappearance of multifinger synergies. In contrast, MU-mode synergies were minimally affected by removing visual feedback off and continued to be robust for the FDS and for the EDC, while being absent for the (FDS + EDC) analysis. We interpret the findings within the theory of hierarchical control of action with spatial referent coordinates. The qualitatively different behavior of the multifinger and MU-mode-based synergies likely reflects the difference in the involved neural circuitry, supraspinal for the former and spinal for the latter.NEW & NOTEWORTHY Two types of synergies, in the space of commands to individual fingers and in the space of motor unit groups, show qualitatively different behaviors during accurate multifinger force-production tasks. After removing visual feedback, finger force synergies disappear, whereas motor unit-based synergies persist. These results point at different neural circuitry involved in these two basic classes of synergies: supraspinal for multieffector synergies, and spinal for motor unit-based synergies.

Intramuscle Synergies: Their Place in the Neural Control Hierarchy.

We accept a definition of synergy introduced by Nikolai Bernstein and develop it for various actions, from those involving the whole body to those involving a single muscle. Furthermore, we use two major theoretical developments in the field of motor control-the idea of hierarchical control with spatial referent coordinates and the uncontrolled manifold hypothesis-to discuss recent studies of synergies within spaces of individual motor units (MUs) recorded within a single muscle. During the accurate finger force production tasks, MUs within hand extrinsic muscles form robust groups, with parallel scaling of the firing frequencies. The loading factors at individual MUs within each of the two main groups link them to the reciprocal and coactivation commands. Furthermore, groups are recruited in a task-specific way with gains that covary to stabilize muscle force. Such force-stabilizing synergies are seen in MUs recorded in the agonist and antagonist muscles but not in the spaces of MUs combined over the two muscles. These observations reflect inherent trade-offs between synergies at different levels of a control hierarchy. MU-based synergies do not show effects of hand dominance, whereas such effects are seen in multifinger synergies. Involuntary, reflex-based, force changes are stabilized by intramuscle synergies but not by multifinger synergies. These observations suggest that multifinger (multimuscle synergies) are based primarily on supraspinal circuitry, whereas intramuscle synergies reflect spinal circuitry. Studies of intra- and multimuscle synergies promise a powerful tool for exploring changes in spinal and supraspinal circuitry across patient populations.

Unintentional drifts in performance during one-hand and two-hand finger force production.

We explored the phenomena of force drifts and unintentional finger force production (enslaving), and their dependence on visual feedback. Predictions have been drawn based on the theory of control with spatial referent coordinates for condition with feedback on instructed (master) finger force, enslaved finger force, and total force for one-hand and two-hand tasks. Subjects produced force under the different feedback conditions without their knowledge. No feedback condition was also used for the one-hand tasks. Overall, feedback of master finger force led to an increase in the enslaved force, feedback on the slave finger force led to a drop in the master force, feedback on the total force led to balanced drifts in the master force down and enslaved force up, and under the no-feedback condition, master and total force drifted down with large variability in the enslaved force drifts. The patterns were the same in both hands in the two-hand tasks when feedback was provided on the forces of one hand only (without subject's knowledge). The index of enslaving always drifted toward higher values. We interpret the findings as reflecting three main factors: drifts in the referent coordinates toward actual finger coordinates, spread of cortical excitation over representations of the fingers, and robust sharing of referent coordinates between the two hands in bimanual tasks. The large consistent drifts in enslaving toward higher values have to be considered in studies of multi-finger synergies.

Intra-muscle Synergies Stabilizing Reflex-mediated Force Changes

We used the framework of the uncontrolled manifold hypothesis to explore force-stabilizing synergies and motor equivalence in the spaces of individual motor unit (MU) firing frequencies. Healthy subjects performed steady force production tasks by pressing with one finger or three fingers of a hand. Surface EMG was used to identify individual MU action potentials. MUs formed stable groups (MU-modes) with parallel scaling of the firing frequency in both flexor digitorum superficialis (FDS) and extensor digitorum communis (EDC) that allowed identifying them with the reciprocal and coactivation commands. Smooth lifting of the fingers by an “inverse piano” device led to an unintentional, reflex-based force increase. There was significantly larger motion in the space of MU-modes that kept the force unchanged (motor equivalent) compared to motion that changed force (non-motor equivalent). The force change was stabilized by co-varying contributions of the MU-modes defined separately for FDS and EDC. In contrast, analysis of the three-finger task in the space of individual finger forces showed no synergies stabilizing total force change. Effects of hand dominance were seen on multi-finger synergies but not intra-muscle synergies. We conclude that spinal mechanisms, such as recurrent inhibition and reflex loops from proprioceptors, contribute significantly to intra-muscle synergies, while multi-finger synergies reflect supra-spinal processes. These results provide methods to explore the contributions of spinal vs supraspinal circuitry to changed motor synergies in populations with a variety of neurological disorders.

Frequency-Dependent Effects on Coordination and Prefrontal Hemodynamics During Finger Force Production Tasks.

Behavioral stability partially depends on the variability of net outcomes by means of the co-varied adjustment of individual elements such as multi-finger forces. The properties of cyclic actions affect stability and variability of the performance as well as the activation of the prefrontal cortex that is an origin of subcortical structure for the coordinative actions. Little research has been done on the issue of the relationship between stability and neuronal response. The purpose of the study was to investigate the changes in the neural response, particularly at the prefrontal cortex, to the frequencies of isometric cyclic finger force production. The main experimental task was to produce finger forces while matching the produced force to sine-wave templates as accurately as possible. Also, the hemodynamics responses of the prefrontal cortex, including oxy-hemoglobin concentration (ΔHbO) and the functional connectivity, were measured using functional near-infrared spectroscopy. The frequency conditions comprised 0.1, 1, and 2 Hz. The uncontrolled manifold (UCM) approach was applied to compute synergy indices in time-series. The relative phase (RP), the coefficient of variation (CV) of the peak and trough force values were computed as the indices of performance accuracy. The statistical parametric mapping (SPM) was implemented to compare the synergy indices of three frequency conditions in time-series. A less accurate performance in the high-frequency condition was caused not by the RP, but mainly by the inconsistent peak force values (CV; p < 0.01, = 0.90). The SPM analysis revealed that the synergy indices were larger in the low-frequency than in high-frequency conditions. Further, the ΔHbO remained unchanged under all frequency conditions, while the functional connectivity decreased with an increase in the frequency of cyclic force production. The current results suggested that the concurrent activation of the prefrontal region mainly depends on the frequency of cyclic force production, which was associated with the strength of stability indices and performance errors. The current study is the first work to uncover the effect of frequency on the multi-finger synergies as to the hemodynamic response in the prefrontal cortex, which possibly provides a clue of the neural mechanism of synergy formation and its changes.

Force-stabilizing synergies can be retained by coordinating sensory-blocked and sensory-intact digits.

The present study examined the effects of selective digital deafferentation on the multi-finger synergies as a function of total force requirement and the number of digits involved in isometric pressing. 12 healthy adults participated in maximal and sub-maximal isometric pressing tasks with or without digital anesthesia to selective digits from the right hand. Our results indicate that selective anesthesia paradigm induces changes in both anesthetized (local) and non-anesthetized (non-local) digits’ performance, including: (1) decreased maximal force abilities in both local and non-local digits; (2) reduced force share during multi-finger tasks from non-local but not local digits; (3) decreased force error-making; and (4) marginally increased motor synergies. These results reinforce the contribution of somatosensory feedback in the process of maximal voluntary contraction force, motor performance, and indicate that somatosensation may play a role in optimizing secondary goals during isometric force production rather than ensuring task performance.

Effects of spastic cerebral palsy on multi-finger coordination during isometric force production tasks.

In this study, we quantified changes in finger interdependence (enslaving), multi-finger synergies, and feedforward modulation of synergy properties (i.e., anticipatory synergy adjustment) during single- and multi-finger force production tasks in individuals with cerebral palsy (CP). Spastic diplegic CP and healthy control subjects performed sets of finger force production tasks by each of the hands, including maximal force production and submaximal quick pulse force production in an isometric condition. The framework of the uncontrolled manifold hypothesis was used to quantify the indices of multi-finger synergies and the anticipatory synergy adjustment (ASA). The CP group showed lower maximal forces and higher indices of finger interdependence (enslaving), while the indices of multi-finger synergies stabilizing total finger forces during stable force production were not different significantly compared to the controls. Further, the time of ASA for the CP group was not delayed. The CP group showed a significantly less drop in the synergy indices during the anticipatory and quick pulse phase compared to the control group, which was accompanied by larger co-contraction indices of the forearm muscles. These findings suggest that the function of assembling motor synergies for stable force production is not affected by CP, while the ability to modulate synergy properties may be impaired with CP partially due to spasticity. The spasticity presumably hampers the purposeful feedforward destabilization of the performance. The results suggest that quantification of multi-digit synergies may provide an alternative tool for quantitative assessment of impaired coordination in the CP individuals.

Age-related changes to motor synergies in multi-joint and multi-finger manipulative skills: a meta-analysis

PurposeThe aim of the current meta-analysis was to examine the extent to which there are differences in upper extremity motor synergies across different age groups in manipulative tasks.MethodsThe studies that used the uncontrolled manifold method to examine the effect of age on motor synergies in multi-joint and multi-finger tasks were selected. Sixteen relevant studies from 1154 articles were selected for the meta-analysis—4 and 12 studies considered multi-joint kinematics and multi-finger kinetic tasks respectively.ResultsThe results of the meta-analysis suggested reduced strength of synergies in multi-finger task in older adults, but this was not the case for synergies in multi-joint task. Part of this age-related difference in finger function is related to the increased variability in total force in grasping tasks. However, reductions in the strength of multi-finger synergies in hand functions following ageing appear to depend on the characteristics of the task.ConclusionsThese findings indicate that the cooperation among fingers to stabilise the total required force to apply for grasping and other fine motor skills is less efficient in older adults that might affect the quality of manipulative tasks.

Effect of Kinetic Degrees of Freedom on Multi-Finger Synergies and Task Performance during Force Production and Release Tasks

Complex structures present in a human body has relatively large degrees-of-freedom (DOFs) as compared to the requirement of a particular task. This phenomenon called motor redundancy initially deemed as a computational problem rather can be understood as having the flexibility to perform the certain task successfully. Hence, the purpose of our study was to examine the positive impact of extra DOFs (redundant DOFs) during force production tasks. For this purpose, an experimental setup was designed to simulate archery-like shooting, and purposeful organization of a redundant set of finger forces determined the stability of important performance variables as well as accurate and precise performance. DOFs were adjusted by changing the number of fingers explicitly involved in the task. The concept of motor synergy and computational framework of uncontrolled manifold (UCM) approach was used to quantify stability indices during finger force production. As a result, accuracy and precision of the task improved with an increase in DOFs. Also, the stability indices of net finger forces and moment increased with active DOFs of fingers. We concluded that the controller actively utilizes extra DOFs to increase the stability of the performance, which is associated with the improved accuracy and precision of the task.

Expectation of movement generates contrasting changes in multifinger synergies in young and older adults.

Anticipatory synergy adjustment (ASA) is a feed-forward control mechanism that describes a continuous decrease in the stability of the current motor state beginning about 150ms prior to a state transition. Recently, we described an associated phenomenon in which the system stability was reduced solely in response to a cue that generates an expectation of a state change, independent of whether the state change actually occurs. Both phenomena are of the same kind (stability reduction), but evoked by distinct antecedent conditions. Since, logically, cuing for movement must occur before the initiation of that movement, we named this new phenomenon 'Stage-1 ASA' and rechristened the well-established version 'Stage-2 ASA'. Here, we used a four-finger, isometric force production task to explore (1) the effect of healthy aging on Stage-1 ASA, and (2) if Stage-1 ASA resulted in a more rapid state change. Young and older adult participants produced 10% of their maximal force when they did not expect to produce any change in the force, and when they expected to change their force in an unknown direction and at an unknown time. In the latter condition, the 10% constant-force phase was followed by a choice reaction time task, in which the participants rapidly changed their force to follow a moving target presented on a computer monitor. Both young and older adults displayed equivalent amount of Stage-1 ASA. This was driven by a 42% reduction in finger-force variability in young adults. In contrast, it was driven by a 38% increase in finger-force variability in older adults. We speculate that the reduction in finger force variability assists the young adults in rapid state changes via two mechanisms: (1) the finger forces occupy a restricted set of states that are optimal for quick state transitions, and (2) lower variability during steady state translates into lower self-motion during state transition. Self-motion is the covariation between finger forces that fails to change the total force. The older adults are unable to adopt this strategy, and the increase in finger-force variability arises from (1) the adoption of an alternative strategy of destabilizing the attractor associated with the current state to facilitate state transitions and (2) the inability to coordinate multiple finger forces. Finally, older adults displayed longer reaction times than young adults, but a clear relation between Stage-1 ASA and consequent behavioral benefit in terms of reduced reaction time remained elusive.

Multi-finger synergies and the muscular apparatus of the hand.

We explored whether the synergic control of the hand during multi-finger force production tasks depends on the hand muscles involved. Healthy subjects performed accurate force production tasks and targeted force pulses while pressing against loops positioned at the level of fingertips, middle phalanges, and proximal phalanges. This varied the involvement of the extrinsic and intrinsic finger flexors. The framework of the uncontrolled manifold (UCM) hypothesis was used to analyze the structure of inter-trial variance, motor equivalence, and anticipatory synergy adjustments prior to the force pulse in the spaces of finger forces and finger modes (hypothetical finger-specific control signals). Subjects showed larger maximal force magnitudes at the proximal site of force production. There were synergies stabilizing total force during steady-state phases across all three sites of force production; no differences were seen across the sites in indices of structure of variance, motor equivalence, or anticipatory synergy adjustments. Indices of variance, which did not affect the task (within the UCM), correlated with motor equivalent motion between the steady states prior to and after the force pulse; in contrast, variance affecting task performance did not correlate with non-motor equivalent motion. The observations are discussed within the framework of hierarchical control with referent coordinates for salient effectors at each level. The findings suggest that multi-finger synergies are defined at the level of abundant transformation between the low-dimensional hand level and higher dimensional finger level while being relatively immune to transformations between the finger level and muscle level. The results also support the scheme of control with two classes of neural variables that define referent coordinates and gains in back-coupling loops between hierarchical control levels.

Stability of hand force production. I. Hand level control variables and multifinger synergies.

We combined the theory of neural control of movement with referent coordinates and the uncontrolled manifold hypothesis to explore synergies stabilizing the hand action in accurate four-finger pressing tasks. In particular, we tested a hypothesis on two classes of synergies, those among the four fingers and those within a pair of control variables, stabilizing hand action under visual feedback and disappearing without visual feedback. Subjects performed four-finger total force and moment production tasks under visual feedback; the feedback was later partially or completely removed. The "inverse piano" device was used to lift and lower the fingers smoothly at the beginning and at the end of each trial. These data were used to compute pairs of hypothetical control variables. Intertrial analysis of variance within the finger force space was used to quantify multifinger synergies stabilizing both force and moment. A data permutation method was used to quantify synergies among control variables. Under visual feedback, synergies in the spaces of finger forces and hypothetical control variables were found to stabilize total force. Without visual feedback, the subjects showed a force drift to lower magnitudes and a moment drift toward pronation. This was accompanied by disappearance of the four-finger synergies and strong attenuation of the control variable synergies. The indexes of the two types of synergies correlated with each other. The findings are interpreted within the scheme with multiple levels of abundant variables.NEW & NOTEWORTHY We extended the idea of hierarchical control with referent spatial coordinates for the effectors and explored two types of synergies stabilizing multifinger force production tasks. We observed synergies among finger forces and synergies between hypothetical control variables that stabilized performance under visual feedback but failed to stabilize it after visual feedback had been removed. Indexes of two types of synergies correlated with each other. The data suggest the existence of multiple mechanisms stabilizing motor actions.

Multi-Finger Interaction and Synergies in Finger Flexion and Extension Force Production.

The aim of this study was to discover finger interaction indices during single-finger ramp tasks and multi-finger coordination during a steady state force production in two directions, flexion, and extension. Furthermore, the indices of anticipatory adjustment of elemental variables (i.e., finger forces) prior to a quick pulse force production were quantified. It is currently unknown whether the organization and anticipatory modulation of stability properties are affected by force directions and strengths of in multi-finger actions. We expected to observe a smaller finger independency and larger indices of multi-finger coordination during extension than during flexion due to both neural and peripheral differences between the finger flexion and extension actions. We also examined the indices of the anticipatory adjustment between different force direction conditions. The anticipatory adjustment could be a neural process, which may be affected by the properties of the muscles and by the direction of the motions. The maximal voluntary contraction (MVC) force was larger for flexion than for extension, which confirmed the fact that the strength of finger flexor muscles (e.g., flexor digitorum profundus) was larger than that of finger extensor (e.g., extensor digitorum). The analysis within the uncontrolled manifold (UCM) hypothesis was used to quantify the motor synergy of elemental variables by decomposing two sources of variances across repetitive trials, which identifies the variances in the uncontrolled manifold (VUCM) and that are orthogonal to the UCM (VORT). The presence of motor synergy and its strength were quantified by the relative amount of VUCM and VORT. The strength of motor synergies at the steady state was larger in the extension condition, which suggests that the stability property (i.e., multi-finger synergies) may be a direction specific quantity. However, the results for the existence of anticipatory adjustment; however, no difference between the directional conditions suggests that feed-forward synergy adjustment (changes in the stability property) may be at least independent of the magnitude of the task-specific apparent performance variables and its direction (e.g., flexion and extension forces).

The synergic control of multi-finger force production: stability of explicit and implicit task components.

Manipulating objects with the hands requires the accurate production of resultant forces including shear forces; effective control of these shear forces also requires the production of internal forces normal to the surface of the object(s) being manipulated. In the present study, we investigated multi-finger synergies stabilizing shear and normal components of force, as well as drifts in both components of force, during isometric pressing tasks requiring a specific magnitude of shear force production. We hypothesized that shear and normal forces would evolve similarly in time and also show similar stability properties as assessed by the decomposition of inter-trial variance within the uncontrolled manifold hypothesis. Healthy subjects were required to accurately produce total shear and total normal forces with four fingers of the hand during a steady-state force task (with and without visual feedback) and a self-paced force pulse task. The two force components showed similar time profiles during both shear force pulse production and unintentional drift induced by turning the visual feedback off. Only the explicitly instructed components of force, however, were stabilized with multi-finger synergies. No force-stabilizing synergies and no anticipatory synergy adjustments were seen for the normal force in shear force production trials. These unexpected qualitative differences in the control of the two force components-which are produced by some of the same muscles and show high degree of temporal coupling-are interpreted within the theory of control with referent coordinates for salient variables. These observations suggest the existence of two classes of neural variables: one that translates into shifts of referent coordinates and defines changes in magnitude of salient variables, and the other controlling gains in back-coupling loops that define stability of the salient variables. Only the former are shared between the explicit and implicit task components.

Changes in Multidigit Synergies and Their Feed-Forward Adjustments in Multiple Sclerosis

ABSTRACTThe authors explored the changes in multidigit synergies in patients with multiple sclerosis (MS) within the framework of the uncontrolled manifold hypothesis. The specific hypotheses were 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 MS patients and age-matched controls (n = 13 in both groups) performed one-finger and multifinger force production tasks involving both accurate steady-state force production and quick force pulses. The patients showed significantly lower maximal finger forces and a tendency toward slower force pulses. Enslaving was increased in MS, but only in the lateral fingers (index and little). Indices of multifinger synergies during steady-state force production were lower in MS, mainly due to the lower amount of intertrial variance that did not affect total force. Anticipatory synergy adjustments were significantly delayed in MS. The results show that MS leads to significant changes in multidigit synergies and feed-forward adjustments of the synergies prior to a quick action. The authors discuss possible contributions of subcortical structures to the impaired synergic control.

On the nature of unintentional action: a study of force/moment drifts during multifinger tasks.

We explored the origins of unintentional changes in performance during accurate force production in isometric conditions seen after turning visual feedback off. The idea of control with referent spatial coordinates suggests that these phenomena could result from drifts of the referent coordinate for the effector. Subjects performed accurate force/moment production tasks by pressing with the fingers of a hand on force sensors. Turning the visual feedback off resulted in slow drifts of both total force and total moment to lower magnitudes of these variables; these drifts were more pronounced in the right hand of the right-handed subjects. Drifts in individual finger forces could be in different direction; in particular, fingers that produced moments of force against the required total moment showed an increase in their forces. The force/moment drift was associated with a drop in the index of synergy stabilizing performance under visual feedback. The drifts in directions that changed performance (non-motor equivalent) and in directions that did not (motor equivalent) were of about the same magnitude. The results suggest that control with referent coordinates is associated with drifts of those referent coordinates toward the corresponding actual coordinates of the hand, a reflection of the natural tendency of physical systems to move toward a minimum of potential energy. The interaction between drifts of the hand referent coordinate and referent orientation leads to counterdirectional drifts in individual finger forces. The results also demonstrate that the sensory information used to create multifinger synergies is necessary for their presence over the task duration.

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.

Dopaminergic modulation of motor coordinaton in Parkinson's disease

BackgroundWe applied the idea of synergies and the framework of the uncontrolled manifold hypothesis to explore the effects of dopamine replacement therapy on finger interaction and coordination in patients with early-stage Parkinson's disease (PD). MethodsEight patients performed single-finger and multi-finger force production tasks with both the dominant and non-dominant hand before (off-drug) and after (on-drug) taking their dopaminergic medications. Synergy indices were defined as co-varied adjustments of commands to fingers that stabilized the total force produced by the hand. ResultsPD patients showed significantly lower maximal finger forces off-drug compared to the on-drug condition, while indices of finger individuation (enslaving) were unchanged. The synergy indices were weaker during steady-state force production off-drug compared to on-drug. Anticipatory adjustments of synergies prior to the quick force pulse initiation were delayed and reduced off-drug as compared to the on-drug condition. These drug effects were observed in both the symptomatic and asymptomatic hands of the patients whose symptoms were limited to one side of the body. ConclusionsThe study demonstrates dopaminergic modulation of motor coordination in PD and supports that the analysis of different components of multi-finger synergies offers a set of indices sensitive to the effects of dopamine replacement therapy in early-stage PD. The results suggest an important role of the basal ganglia in synergy formation and in feed-forward synergy adjustments. Future studies using these methods may yield more objective, quantitative biomarker(s) of motor coordination impairments in PD, and better understanding of subcortical involvement in the neural control of natural actions.

Effects of Parkinson’s disease on optimization and structure of variance in multi-finger tasks

We explored the role of the basal ganglia in two components of multi-finger synergies by testing a group of patients with early-stage Parkinson's disease and a group of healthy controls. Synergies were defined as co-varied adjustments of commands to individual fingers that reduced variance of the total force and moment of force. The framework of the uncontrolled manifold hypothesis was used to quantify such co-variation patterns, while average performance across repetitive trials (sharing patterns) was analyzed using the analytical inverse optimization (ANIO) approach. The subjects performed four-finger pressing tasks that involved the accurate production of combinations of the total force and total moment of force and also repetitive trials at two selected combinations of the total force and moment. The ANIO approach revealed significantly larger deviations of the experimental data planes from an optimal plane for the patients compared to the control subjects. The synergy indices computed for total force stabilization were significantly higher in the control subjects compared to the patients; this was not true for synergy indices computed for moment of force stabilization. The differences in the synergy indices were due to the larger amount of variance that affected total force in the patients, while the amount of variance that did not affect total force was comparable between the groups. We conclude that the basal ganglia play an important role in both components of synergies reflecting optimization of the sharing patterns and stability of performance with respect to functionally important variables.

Effects of muscle vibration on multi-finger interaction and coordination

The purpose of this study was to investigate the effects of changes in the proprioceptive signals induced by muscle vibration on multi-finger interaction and coordination. We hypothesized that unintended force production by non-instructed fingers (enslaving) would increase with muscle vibration while synergy indices during steady-state force production would drop. The framework of the uncontrolled manifold hypothesis was used to quantify indices of multi-finger synergies stabilizing total force during steady-state force production and anticipatory changes in these indices (anticipatory synergy adjustments, ASAs) in preparation to a quick force pulse production with and without hand-muscle vibration at 80 Hz. The dominant hands of twelve healthy right-handed subjects were tested under three conditions: no vibration, vibration of the palmar surface of the hand, and vibration of the forearm applied over the flexor muscles. There were no significant effects of vibration on maximal voluntary force. The magnitude of enslaving was larger during vibration of the hand compared to the other two conditions. During steady-state force production, strong synergies stabilizing total force were seen in all three conditions; however, indices of force-stabilizing synergies were lower during vibration of the hand. Prior to the force pulse initiation, the synergy index started to drop earlier and over a larger magnitude without vibration compared to either vibration condition. Effects of vibration on enslaving and synergy index may be due to diffuse reflex effects of the induced afferent activity on alpha-motoneuronal pools innervating the extrinsic flexor compartments. We conclude that multi-finger synergies are not based on signals from muscle receptors. The smaller synergy indices and ASAs may reflect supraspinal effects of the vibration-induced afferent activity, in particular its interactions with trans-thalamic loops.