Motor Equivalence Research Articles

Motor imagery from brain to muscle: a commentary on Bach et al., (2022).

In a recent article entitled "Why motor imagery is not really motoric: towards a re-conceptualization in terms of effect-based action control", Bach et al. nicely renewed the concept of motor equivalence between actual movement and motor imagery (MI), i.e. the mental simulation of an action without its corresponding motor output. Their approach is largely based on behavioral studies and, to a lesser extent, on the literature using cerebral imagery. However, the literature on cortico-spinal circuitry modulation during MI can provide further, interesting aspects. Indeed, when it comes to addressing the motor system, one should consider the whole path from brain region to muscle contraction, including sub-cortical structures such as the spinal circuitry. This commentary aims at bridging this gap by providing supplemental evidence and outlining a complementary approach.

Skydiving technique analysis from a control engineering perspective: Developing a tool for studying human motor equivalence

This study offers a novel mathematical approach to sports technique analysis and motor equivalency investigation. The method is aimed to deal with intensive interaction between the environment and a trainee, and activities for which imitation learning is less efficient and movements must be established individually. The free-fall stage of skydiving is investigated, when aerial manoeuvres are performed by changing the body posture and thus deflecting the surrounding airflow. The natural learning process of body flight is hard and protracted as the required movements are not similar to our natural movement repertoire, are often counter-intuitive, and highly dependant on anthropometric factors. The proposed method extracts the trainee’s movement patterns via Principal Component Analysis, and analyses the performed manoeuvres by modelling the dynamic response of the closed-loop feedback control system comprising the body actuated by each pattern and the environment. The method can provide a valuable insight into the subject’s learning process. The main novelty is that instead of comparing trainee’s movements to a template or a movement pattern, extracted from a top-rated athlete, we offer an independent way of technique analysis. Design tools of automatic control theory are utilised to suggest technique modifications that provide the desired performance improvement. An unconventional insight into the motor equivalence problem is achieved: Multiple body degrees-of-freedom are needed for shaping dynamic characteristics of the controlled plant. Movement patterns synergies provide a trade-off between these characteristics, e.g. stability and agility. The method is demonstrated on three case studies of aerial rotation of skilled, less-skilled, and elite skydivers.

The effect of the preferred hand on drawing movement

The observation that different effectors can execute the same movement suggests functional equivalences driven by limb independent representation of action in the central nervous system. A common invariant motor behavior is the speed and curvature coupling (the 1/3 power law), a low dimensional (abstract) descriptor of movement which is resilient to different sensorimotor contexts. Our purpose is to verify the consistency of such motor equivalence during a drawing task, by testing the effect of manual dominance and movement speed on motor performance. We hypothesize that abstract kinematic variables are not the most resistant to speed or limb effector changes. The results show specific effects of speed and hand side on the drawing task. Movement duration, speed-curvature covariation, and maximum velocity were not significantly affected by hand side, while geometrical features were strongly speed and limb dependent. However, intra-trial analysis performed over the successive drawing movements reveals a significant hand side effect on the variability of movement vigor and velocity-curvature relationship (the 1/3 PL). The identified effects of speed and hand dominance on the kinematic parameters suggest different neural strategies, in a pattern that does not go from the most abstract to the least abstract component, as proposed by the traditional hierarchical organization of the motor plan.

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.

Turning a blind eye to motor differences leads to bias in estimating action-related auditory ERP attenuation

Event-related potential (ERP) studies investigating the processing of self-induced stimuli often rely on the assumption that ballistic actions and motor ERPs are constant across different sets of action effects. Since recent studies challenge this motor equivalence assumption, we examined whether neglecting effect-related motor differences can bias the estimation of auditory ERPs in a typical action-related ERP attenuation paradigm. We increased action variability with a force production task and selected an event subset in which the motor equivalence assumption was true. ERP attenuation estimated in this subset was compared with attenuation obtained in the standard task, where motor differences were not controlled. Violation of the motor equivalence assumption resulted in a positive deflection overlapping auditory ERPs elicited by self-induced sounds, leading to the overestimation of N1- and underestimation of P2-attenuation. This demonstrates that sensory-effect-related motor differences should be considered when separating sensory and motor components in ERPs elicited by self-induced stimuli.

LaDIVA: A neurocomputational model providing laryngeal motor control for speech acquisition and production.

Many voice disorders are the result of intricate neural and/or biomechanical impairments that are poorly understood. The limited knowledge of their etiological and pathophysiological mechanisms hampers effective clinical management. Behavioral studies have been used concurrently with computational models to better understand typical and pathological laryngeal motor control. Thus far, however, a unified computational framework that quantitatively integrates physiologically relevant models of phonation with the neural control of speech has not been developed. Here, we introduce LaDIVA, a novel neurocomputational model with physiologically based laryngeal motor control. We combined the DIVA model (an established neural network model of speech motor control) with the extended body-cover model (a physics-based vocal fold model). The resulting integrated model, LaDIVA, was validated by comparing its model simulations with behavioral responses to perturbations of auditory vocal fundamental frequency (fo) feedback in adults with typical speech. LaDIVA demonstrated capability to simulate different modes of laryngeal motor control, ranging from short-term (i.e., reflexive) and long-term (i.e., adaptive) auditory feedback paradigms, to generating prosodic contours in speech. Simulations showed that LaDIVA's laryngeal motor control displays properties of motor equivalence, i.e., LaDIVA could robustly generate compensatory responses to reflexive vocal fo perturbations with varying initial laryngeal muscle activation levels leading to the same output. The model can also generate prosodic contours for studying laryngeal motor control in running speech. LaDIVA can expand the understanding of the physiology of human phonation to enable, for the first time, the investigation of causal effects of neural motor control in the fine structure of the vocal signal.

Neural representation in M1 and S1 cortex of bilateral hand actions during prehension.

Bimanual movements that require coordinated actions of the two hands may be coordinated by synchronous bilateral activation of somatosensory and motor cortical areas in both hemispheres, by enhanced activation of individual neurons specialized for bimanual actions, or by both mechanisms. To investigate cortical neural mechanisms that mediate unimanual and bimanual prehension, we compared actions of the left and right hands in a reach to grasp-and-pull instructed-delay task. Spike trains were recorded with multiple electrode arrays placed in the hand area of primary motor (M1) and somatosensory (S1) cortex of the right hemisphere in macaques, allowing us to measure and compare the relative timing, amplitude, and synchronization of cortical activity in these areas as animals grasped and manipulated objects that differed in shape and location. We report that neurons in the right hemisphere show common task-related firing patterns for the two hands but actions of the ipsilateral hand elicited weaker and shorter-duration responses than those of the contralateral hand. We report significant bimanual activation of neurons in M1 but not in S1 cortex when animals have free choice of hand use in prehension tasks. Population ensemble responses in M1 thereby provide an accurate depiction of hand actions during skilled manual tasks. These studies also demonstrate that somatosensory cortical areas serve important cognitive and motor functions in skilled hand actions. Bilateral representation of hand actions may serve an important role in "motor equivalence" when the same movements are performed by either hand and in transfer of skill learning between the hands.NEW & NOTEWORTHY Humans can manipulate small objects with the right or left hand but typically select the dominant hand to handle them. We trained monkeys to grasp and manipulate objects with either hand, while recording neural activity in primary motor (M1) and somatosensory (S1) cortex. Actions of both hands activate M1 neurons, but S1 neurons respond only to the contralateral hand. Bilateral sensitivity in M1 may aid skill transfer between hands after stroke or head injury.

A Vexing Question in Motor Control: The Degrees of Freedom Problem.

The human “marionette” is extremely complex and multi-articulated: anatomical redundancy (in terms of Degrees of Freedom: DoFs), kinematic redundancy (movements can have different trajectories, velocities, and accelerations and yet achieve the same goal, according to the principle of Motor Equivalence), and neurophysiological redundancy (many more muscles than DoFs and multiple motor units for each muscle). Although it is quite obvious that such abundance is not noxious at all because, in contrast, it is instrumental for motor learning, allowing the nervous system to “explore” the space of feasible actions before settling on an elegant and possibly optimal solution, the crucial question then boils down to figure out how the nervous system “chooses/selects/recruits/modulates” task-dependent subsets of countless assemblies of DoFs as functional motor synergies. Despite this daunting conceptual riddle, human purposive behavior in daily life activities is a proof of concept that solutions can be found easily and quickly by the embodied brain of the human cognitive agent. The point of view suggested in this essay is to frame the question above in the old-fashioned but still seminal observation by Marr and Poggio that cognitive agents should be regarded as Generalized Information Processing Systems (GIPS) and should be investigated according to three nearly independent but complementary levels of analysis: 1) the computational level, 2) the algorithmic level, and 3) the implementation level. In this framework, we attempt to discriminate as well as aggregate the different hypotheses and solutions proposed so far: the optimal control hypothesis, the muscle synergy hypothesis, the equilibrium point hypothesis, or the uncontrolled manifold hypothesis, to mention the most popular ones. The proposed GIPS follows the strategy of factoring out shaping and timing by adopting a force-field based approach (the Passive Motion Paradigm) that is inspired by the Equilibrium Point Hypothesis, extended in such a way to represent covert as well overt actions. In particular, it is shown how this approach can explain spatio-temporal invariances and, at the same time, solve the Degrees of Freedom Problem.

Sex-specific effects of a repetitive fatiguing task on stability: Analysis with motor equivalence model

Although studies showed that several internal factors affect task-specific stability, the sex-specific effects of fatigue on whole-limb stability during a semi-cycle repetitive pointing task remain unidentified. Synergy and Motor Equivalence concepts in the UCM framework have been developed to explain task-specific stability. The motor equivalence model quantifies the amount of deviation in the space of elemental variables that occurs in two directions; one that preserves the performance variable (good variance), and the other that affects it (bad variance). Synergy index (the difference between good and bad variance divided by the total variance > 0) represent stability in performing a task. Healthy adults (n = 26, 13F; age: 35.3 ± 10.6 yrs.) performed an RPT by moving their dominant arm between a proximal target and a distal target in a standing position until near fatigue (Borg CR10 rating 8/10). Tridimensional kinematics of trunk, upper arm, forearm, and hand segments were captured by high-resolution cameras every minute, and joint angles were extracted according to the ZX′Y″ Euler sequence. Results showed the synergy > 0 for both women and men, reflecting synergies stabilizing the endpoint coordinate in both Non-Fatigue and Fatigue conditions. Statistics (ANOVA) showed a significant Condition * Sex effect (p = 0.01), with higher good (by 0.19 ± 0.1 rad) and bad variances (by 0.15 ± 0.09 rad) in women compared to men after fatigue. Higher good and bad variability, with no change in women's performance could represent a less stable strategy, leading to the development of risk factors for neck-shoulder disorders.

Synergic control of action in levodopa-naïve Parkinson's disease patients: II. Multi-muscle synergies stabilizing vertical posture.

Postural instability is a major disabling feature in Parkinson's disease (PD). We quantified the organization of leg and trunk muscles into synergies stabilizing the center of pressure (COP) coordinate within the uncontrolled manifold hypothesis in levodopa-naïve patients with PD and age-matched control subjects. The main hypothesis was that changes in the synergic control of posture are present early in the PD process even before levodopa exposure. Eleven levodopa-naïve patients with PD and 11 healthy controls performed whole-body cyclical voluntary sway tasks and a self-initiated load-release task during standing on a force plate. Surface electromyographic activity in 13 muscles on the right side of the body was analyzed to identify muscle groups with parallel scaling of activation levels (M-modes). Data were collected both before ("off-drug") and approximately 60min after the first dose of 25/100 carbidopa/levodopa ("on-drug"). COP-stabilizing synergies were quantified for the load-release task. Levodopa-naïve patients with PD showed no COP-stabilizing synergy "off-drug", whereas controls showed posture-stabilizing multi-M-mode synergy. "On-drug", patients with PD demonstrated a significant increase in the synergy index. There were no significant drug effects on the M-mode composition, anticipatory postural adjustments, indices of motor equivalence, or indices of COP variability. The results suggest that levodopa-naïve patients with PD already show impaired posture-stabilizing multi-muscle synergies that may be used as promising behavioral biomarkers for emerging postural disorders in PD. Moreover, levodopa modified synergy metrics differently in these levodopa-naïve patients compared to a previous study of patients on chronic antiparkinsonian medications (Falaki et al. in J Electromyogr Kinesiol 33:20-26, 2017a), suggesting different neurocircuitry involvement.

From motor to visually guided bimanual affordance learning

The mechanisms of how the brain orchestrates multi-limb joint action have yet to be elucidated and few computational sensorimotor (SM) learning approaches have dealt with the problem of acquiring bimanual affordances. We propose a series of bidirectional (forward/inverse) SM maps and its associated learning processes that generalize from uni- to bimanual interaction (and affordances) naturally, reinforcing the motor equivalence property. The SM maps range from a SM nature to a solely sensory one: full body control, delta SM control (through small action changes), delta sensory co-variation (how body-related perceptual cues covariate with object-related ones). We make several contributions on how these SM maps are learned: (1) Context and Behavior-Based Babbling: generalizing goal babbling to the interleaving of absolute and local goals including guidance of reflexive behaviors; (2) Event-Based Learning: learning steps are driven by visual, haptic events; and (3) Affordance Gradients: the vectorial field gradients in which an object can be manipulated. Our modeling of bimanual affordances is in line with current robotic research in forward visuomotor mappings and visual servoing, enforces the motor equivalence property, and is also consistent with neurophysiological findings like the multiplicative encoding scheme.

Preparation to a quick whole-body action: control with referent body orientation and multi-muscle synergies.

We examined the control of postural stability in preparation to a discrete, quick whole-body sway toward a target and back to the initial position. Several predictions were tested based on the theory of control with referent body orientation and the notion of multi-muscle synergies stabilizing center of pressure (COP) coordinate. Healthy, young adults performed fast, discrete whole-body motion forward-and-back and backward-and-back under visual feedback on the COP. We used two methods to assess COP stability, analysis of inter-trial variance and analysis of motor equivalence in the muscle activation space. Actions were always preceded by COP counter-movements. Backward COP shifts were faster, and the indices of multi-muscle synergies stabilizing COP were higher prior to those actions. Patterns of muscle activation at the motion onset supported the idea of a gradual shift in the referent body orientation. Prior to the backward movements, there was a trend toward higher muscle co-activation, compared to reciprocal activation. We found strong correlations between the sets of indices of motor equivalence and those of inter-trial variance. Overall, the results support the theory of control with referent coordinates and the idea of multi-muscle synergies stabilizing posture by confirming a number of non-trivial predictions based on these concepts. The findings favor using indices of motor equivalence in clinical studies to minimize the number of trials performed by each subject.

A process account of the uncontrolled manifold structure of joint space variance in pointing movements

In many situations, the human movement system has more degrees of freedom than needed to achieve a given movement task. Martin et al. (Neural Comput 21(5):1371–1414, 2009) accounted for signatures of such redundancy like self-motion and motor equivalence in a process model in which a neural oscillator generated timed end-effector virtual trajectories that a neural dynamics transformed into joint virtual trajectories while decoupling task-relevant and task-irrelevant combinations of joint angles. Neural control of muscle activation and the biomechanical dynamics of the arm were taken into account. The model did not address the main signature of redundancy, however, the UCM structure of variance: Many experimental studies have shown that across repetitions, variance of joint configuration trajectories is structured. Combinations of joint angles that affect task variables (lying in the uncontrolled manifold, UCM) are much more variable than combinations of joint angles that do not. This finding has been robust across movement systems, age, and tasks and is often preserved in clinical populations as well. Here, we provide an account for the UCM structure of variance by adding four types of noise sources to the model of Martin et al. (Neural Comput 21(5):1371–1414, 2009). Comparing the model to human pointing movements and systematically examining the role of each noise source and mechanism, we identify three causes of the UCM effect, all of which, we argue, contribute: (1) the decoupling of motor commands across the task-relevant and task-irrelevant subspaces together with “neural” noise at the level of these motor commands; (2) “muscle noise” combined with imperfect control of the limb; (3) back-coupling of sensed joint configurations into the motor commands which then yield to the sensed joint configuration within the UCM.

IDeLog: Iterative Dual Spatial and Kinematic Extraction of Sigma-Lognormal Parameters.

The Kinematic Theory of rapid movements and its associated Sigma-Lognormal model have been extensively used in a large variety of applications. While the physical and biological meaning of the model have been widely tested and validated for rapid movements, some shortcomings have been detected when it is used with continuous long and complex movements. To alleviate such drawbacks, and inspired by the motor equivalence theory and a conceivable visual feedback, this paper proposes a novel framework to extract the Sigma-Lognormal parameters, namely iDeLog. Specifically, iDeLog consists of two steps. The first one, influenced by the motor equivalence model, separately derives an initial action plan defined by a set of virtual points and angles from the trajectory and a sequence of lognormals from the velocity. In the second step, based on a hypothetical visual feedback compatible with an open-loop motor control, the virtual target points of the action plan are iteratively moved to improve the matching between the observed and reconstructed trajectory and velocity. During experiments conducted with handwritten signatures, iDeLog obtained promising results as compared to the previous development of the Sigma-Lognormal.

Individual preferences in motor coordination seen across the two hands: relations to movement stability and optimality.

The framework of the uncontrolled manifold (UCM) hypothesis was used to explore variables related to stability of task performance in the two hands of young healthy individuals. Fourteen young adults performed four-finger accurate constant force production tasks interrupted by a voluntary quick force pulse production and by an externally imposed displacement of all fingers. Three groups of variables were used to quantify stability of steady force production: (1) indices of the inter-trial variance were computed within the UCM and orthogonal to the UCM; (2) indices of motor equivalence were computed between steady-state intervals separated by the force pulse and by thefinger-lifting episode; and (3) referent coordinate and apparent stiffness were computed using the data during the ascending phase of the finger-lifting episode. In another task, the subjects performed accurate constant force production with visual feedback removal after the 8th second, and the drop in the total force after the removal was computed. There were differences between the right and left hand in some outcome variables such as variance within the UCM, and the timing of anticipatory synergy adjustments prior to the force pulse, consistent with the dynamic dominance hypothesis. There were significant correlations between the two hands for indices that were unrelated to accuracy of performance: variance within the UCM, index of motor equivalence, referent coordinate, apparent stiffness, and the drop of total force after visual feedback removal. We interpret these findings within the concept of stability-optimality trade-off. In particular, we conclude that individual subjects select particular, person-specific solutions within the spectrum allowed by the explicit task constraints, and this choice is consistent between the two hands. We conclude with a hypothesis that selecting specific solutions within the stability-optimality trade-off may represent an individual's personal preference consistent between the two hands.

Stability of steady hand force production explored across spaces and methods of analysis.

We used the framework of the uncontrolled manifold (UCM) hypothesis and explored the reliability of several outcome variables across different spaces of analysis during a very simple four-finger accurate force production task. Fourteen healthy, young adults performed the accurate force production task with each hand on 3 days. Small spatial finger perturbations were generated by the "inverse piano" device three times per trial (lifting the fingers 1cm/0.5s and lowering them). The data were analyzed using the following main methods: (1) computation of indices of the structure of inter-trial variance and motor equivalence in the space of finger forces and finger modes, and (2) analysis of referent coordinates and apparent stiffness values for the hand. Maximal voluntary force and the index of enslaving (unintentional finger force production) showed good to excellent reliability. Strong synergies stabilizing total force were reflected in both structure of variance and motor equivalence indices. Variance within the UCM and the index of motor equivalent motion dropped over the trial duration and showed good to excellent reliability. Variance orthogonal to the UCM and the index of non-motor equivalent motion dropped over the 3 days and showed poor to moderate reliability. Referent coordinate and apparent stiffness indices co-varied strongly and both showed good reliability. In contrast, the computed index of force stabilization showed poor reliability. The findings are interpreted within the scheme of neural control with referent coordinates involving the hierarchy of two basic commands, the r-command and c-command. The data suggest natural drifts in the finger force space, particularly within the UCM. We interpret these drifts as reflections of a trade-off between stability and optimization of action. The implications of these findings for the UCM framework and future clinical applications are explored in the discussion. Indices of the structure of variance and motor equivalence show good reliability and can be recommended for applied studies.

Stability of vertical posture explored with unexpected mechanical perturbations: synergy indices and motor equivalence.

We explored the relations between indices of mechanical stability of vertical posture and synergy indices under unexpected perturbations. The main hypotheses predicted higher posture-stabilizing synergy indices and higher mechanical indices of center of pressure stability during perturbations perceived by subjects as less challenging. Healthy subjects stood on a force platform and held in fully extended arms a bar attached to two loads acting downward and upward. One of the loads was unexpectedly released by the experimenter causing a postural perturbations. In different series, subjects either knew or did not know which of the two loads would be released. Forward perturbations were perceived as more challenging and accompanied by co-activation patterns among the main agonist-antagonist pairs. Backward perturbation led to reciprocal muscle activation patterns and was accompanied by indices of mechanical stability and of posture-stabilizing synergy which indicated higher stability. Changes in synergy indices were observed as early as 50-100ms following the perturbation reflecting involuntary mechanisms. In contrast, predictability of perturbation direction had weak or no effect on mechanical and synergy indices of stability. These observations are interpreted within a hierarchical scheme of synergic control of motor tasks and a hypothesis on the control of movements with shifts of referent coordinates. The findings show direct correspondence between stability indices based on mechanics and on the analysis of multi-muscle synergies. They suggest that involuntary posture-stabilizing mechanisms show synergic organization. They also show that predictability of perturbation direction has strong effects on anticipatory postural adjustment but not corrective adjustments. We offer an interpretation of co-activation patterns that questions their contribution to postural stability.

Response properties of saccade-related neurons of the post-arcuate premotor cortex.

We studied the phasic saccade-related discharges of single neurons (S neurons) of the premotor cortex of female rhesus monkeys, mostly in the caudal bank of the arcuate sulcus. As described in previous work from our laboratory (Neromyliotis E, Moschovakis AK. Front Behav Neurosci 11: 1-21, 2017), some of these cells emitted phasic discharges for coordinated movements of the eyes and hand as well as for movements of either effector executed in isolation (motor equivalence, Meq). Other cells (S) did not emit phasic discharges for hand movements unaccompanied by saccades. In contrast to frontal eye field (FEF) neurons, but similar to forelimb-related neurons (H neurons) and Meq cells, the discharges of S cells did not display contralateral bias; their on-directions were as likely to be ipsiversive as contraversive. Because the onset of their discharge preceded that of FEF neurons, S cells are unlikely to convey to their targets corollary discharges of the FEF. We also encountered a small number of neurons that could function as logic gates: cells that discharged for saccades if they were not accompanied by hand movements, cells that discharged for saccades or movements of the hand but not for coordinated movements of both effectors, and cells that discharged only for coordinated movements of the eyes and the hand but not when one of the effectors moved unaccompanied by the other. Our findings are discussed in terms of sequences of decision processes stitching effector-specific motor plans onto effector-invariant movement primitives. NEW & NOTEWORTHY The premotor cortex, traditionally associated with skeletomotor control, is shown to contain cells that emit strong discharges time-linked to saccades but not for hand movements unaccompanied by saccades (S cells). Unlike frontal eye field (FEF) neurons, the S cells of the premotor cortex did not display contralateral bias, and because their presaccadic discharges preceded those of FEF neurons, they are unlikely to serve as conveyors of FEF efferent discharges.

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.

Reduced Kinematic Redundancy and Motor Equivalence During Whole-Body Reaching in Individuals With Chronic Stroke

Kinematic redundancy of the human body provides abundant movement patterns to accomplish the same motor goals (motor equivalence). Compensatory movement patterns such as excessive trunk displacement in stroke subjects during reaching can be viewed as a consequence of the motor equivalent process to accomplish a task despite limited available ranges in some joints. However, despite compensations, the ability to adapt reaching performance when perturbations occur may still be limited when condition-specific changes of joint angles are required. We addressed this hypothesis in individuals with and without stroke for reaching a target placed beyond arm reach in standing while flexing the hips (free-hip condition). In randomly selected trials, hip flexion was unexpectedly blocked, forcing subjects to take a step (blocked-hip condition). In additional trials, subjects took an intentional step while reaching the target (intentional-step condition). In blocked-hip trials, healthy subjects maintained smooth and precise endpoint trajectories by adapting temporal and spatial interjoint coordination to neutralize the effect of the perturbation. However, the ability to produce motor equivalent solutions was reduced in subjects with stroke, evidenced by substantial overshoot errors in endpoint position, reduced movement smoothness and less adaptive elbow-shoulder interjoint coordination. Movement adaptability was more limited in stroke subjects who used more compensatory movements for unperturbed reaching. Results suggest that subjects with mild-to-moderate stroke only partially adapted arm joint movements to maintain reaching performance. Therapeutic efforts to enhance the ability of individuals with stroke to find a larger number of task-relevant motor solutions (adaptability) may improve upper limb recovery.