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The goal of the research is to reconstruct the unknown cost (objective) function(s) presumably used by the neural controller for sharing the total force among individual fingers in multifinger prehension. The cost function was determined from experimental data by applying the recently developed Analytical Inverse Optimization (ANIO) method (Terekhov et al. 2010). The core of the ANIO method is the Theorem of Uniqueness that specifies conditions for unique (with some restrictions) estimation of the objective functions. In the experiment, subjects (n = 8) grasped an instrumented handle and maintained it at rest in the air with various external torques, loads, and target grasping forces applied to the object. The experimental data recorded from 80 trials showed a tendency to lie on a 2-dimensional hyperplane in the 4-dimensional finger-force space. Because the constraints in each trial were different, such a propensity is a manifestation of a neural mechanism (not the task mechanics). In agreement with the Lagrange principle for the inverse optimization, the plane of experimental observations was close to the plane resulting from the direct optimization. The latter plane was determined using the ANIO method. The unknown cost function was reconstructed successfully for each performer, as well as for the group data. The cost functions were found to be quadratic with nonzero linear terms. The cost functions obtained with the ANIO method yielded more accurate results than other optimization methods. The ANIO method has an evident potential for addressing the problem of optimization in motor control. [ABSTRACT FROM AUTHOR]

The method of multidimensional scaling was applied to matrices of finger interaction (IFM) computed for individual participants for finger force production tasks. When IFMs for young controls, elderly, and persons with Down syndrome were pooled, only two dimensions described interpersonal differences; these were related to total force and to the total amount of enslaving. When IFMs for each group were analyzed separately, subpopulation-specific dimensions were found. Potentially, this analysis can be applied to discover meaningful dimensions that reflect differences in indices of finger interaction across and within subpopulations which differ in their apparent ability to use the hand. It may also be useful for tracking changes in finger interaction that occur in the process of specialized training or motor rehabilitation. [ABSTRACT FROM AUTHOR]

We address the nature of unintentional changes in performance in two papers. This second paper tested hypotheses related to stability of task-specific performance variables estimated using the framework of the uncontrolled manifold (UCM) hypothesis. Our first hypothesis was that selective stability of performance variables would be observed even when the magnitudes of those variables drifted unintentionally because of the lack of visual feedback. Our second hypothesis was that stability of a variable would depend on the number of explicit task constraints. Subjects performed four-finger isometric pressing tasks that required the accurate production of a combination of total moment and total force with natural or modified finger involvement under full visual feedback, which was removed later for some or all of the salient variables. We used inter-trial analysis of variance and drifts in the space of finger forces within the UCM and within the orthogonal to the UCM space. The two variance components were used to estimate a synergy index stabilizing the force/moment combination, while the two drift components were used to estimate motor equivalent and non-motor equivalent force changes, respectively. Without visual feedback, both force and moment drifted toward lower absolute magnitudes. The non-motor equivalent component of motion in the finger force space was larger than the motor equivalent component for variables that stopped receiving visual feedback. In contrast, variables that continued to receive visual feedback showed larger motor equivalent component, compared to non-motor equivalent component, over the same time interval. These data falsified the first hypothesis; indeed, selective stabilization of a variable over the duration of a trial allows expecting comparably large motor equivalent components both with and without visual feedback. Adding a new constraint (presented as a target magnitude of middle finger force) resulted in a drop in the synergy index in support of the second hypothesis. We interpret the force drift as a natural relaxation process toward states with lower potential energy in the physical (physiological) system involved in the task. The results show that presenting sensory feedback on a performance variable makes synergies stabilizing that variable dependent on that particular sensory feedback.

We address the nature of unintentional changes in performance in two papers. This first paper tested a hypothesis that unintentional changes in performance variables during continuous tasks without visual feedback are due to two processes. First, there is a drift of the referent coordinate for the salient performance variable toward the actual coordinate of the effector. Second, there is a drift toward minimum of a cost function. We tested this hypothesis in four-finger isometric pressing tasks that required the accurate production of a combination of total moment and total force with natural and modified finger involvement. Subjects performed accurate force-moment production tasks under visual feedback, and then visual feedback was removed for some or all of the salient variables. Analytical inverse optimization was used to compute a cost function. Without visual feedback, both force and moment drifted slowly toward lower absolute magnitudes. Over 15 s, the force drop could reach 20% of its initial magnitude while moment drop could reach 30% of its initial magnitude. Individual finger forces could show drifts toward both higher and lower forces. The cost function estimated using the analytical inverse optimization reduced its value as a consequence of the drift. We interpret the results within the framework of hierarchical control with referent spatial coordinates for salient variables at each level of the hierarchy combined with synergic control of salient variables. The force drift is discussed as a natural relaxation process toward states with lower potential energy in the physical (physiological) system involved in the task.

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.

We applied the theory of synergies to analyze the processes that lead to unintentional decline in isometric fingertip force when visual feedback of the produced force is removed. We tracked the changes in hypothetical control variables involved in single fingertip force production based on the equilibrium-point hypothesis, namely the fingertip referent coordinate (R FT ) and its apparent stiffness (C FT ). The system's state is defined by a point in the {R FT ; C FT } space. We tested the hypothesis that, after visual feedback removal, this point (1) moves along directions leading to drop in the output fingertip force, and (2) has even greater motion along directions that leaves the force unchanged. Subjects produced a prescribed fingertip force using visual feedback and attempted to maintain this force for 15 s after the feedback was removed. We used the "inverse piano" apparatus to apply small and smooth positional perturbations to fingers at various times after visual feedback removal. The time courses of R FT and C FT showed that force drop was mostly due to a drift in R FT toward the actual fingertip position. Three analysis techniques, namely hyperbolic regression, surrogate data analysis, and computation of motor-equivalent and non-motor-equivalent motions, suggested strong covariation in R FT and C FT stabilizing the force magnitude. Finally, the changes in the two hypothetical control variables {R FT ; C FT } relative to their average trends also displayed covariation. On the whole, the findings suggest that unintentional force drop is associated with (a) a slow drift of the referent coordinate that pulls the system toward a low-energy state and (b) a faster synergic motion of R FT and C FT that tends to stabilize the output fingertip force about the slowly drifting equilibrium point.

The authors explored application of analytical inverse optimization (ANIO) method to the normal finger forces in unimanual and bimanual prehensile tasks with discrete and continuously changing constraints. The subjects held an instrumented handle vertically with one or two hands. The external torque and grip force changed across trials or within a trial continuously. Principal component analysis showed similar percentages of variance accounted for by the first two principal components across tasks and conditions. Compared to unimanual tasks, bimanual tasks showed significantly more frequent inability to find a cost function leading to a stable solution. In cases of stable solutions, similar second-order polynomials were computed as cost functions across tasks and condition. The bimanual tasks, however, showed significantly worse goodness-of-fit index values. The authors show that ANIO can be used in tasks with slowly changing constraints making it an attractive tool to study optimality of performance in special populations. They also show that ANIO can fail in multifinger tasks, likely due to irreproducible behavior across trials, more likely to happen in bimanual tasks compared to unimanual tasks.

We investigated multidigit synergies stabilizing components of the resultant force vector during joint performance of a static prehension task by two persons as compared to similar tasks performed by a single person using both hands. Subjects transferred the instrumented handle from the right hand to the left hand (one-person condition) or passed that handle to another person (two-person condition) while keeping the handle's position and orientation stationary. Only three digits were involved per hand, the thumb, the index finger, and the middle finger; the forces and moments produced by the digits were measured by six-component sensors. We estimated the performance-stabilizing synergies within the uncontrolled manifold framework by quantifying the intertrial variance structure of digit forces and moments. The analysis was performed at three levels: between hands, between virtual finger and virtual thumb (imagined digits producing the same mechanical variables as the corresponding actual digits combined) produced by the two hands (in both interpersonal and intrapersonal conditions), and between the thumb and virtual finger for one hand only. Additionally, we performed correlation and phase synchronization analyses of resultant tangential forces and internal normal forces. Overall, the one-person conditions were characterized by higher amount of intertrial variance that did not affect resultant normal force components, higher internal components of normal forces, and stronger synchronization of the normal forces generated by the hands. Our observations suggest that in two-person tasks, when participants try to achieve a common mechanical outcome, the performance-stabilizing synergies depend on non-visual information exchange, possibly via the haptic and proprioceptive systems. Therefore, synergies quantified in tasks using visual feedback only may not be generalizable to more natural 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., (Copyright © 2016 the American Physiological Society.)

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., (Copyright © 2015 IBRO. Published by Elsevier Ltd. All rights reserved.)

We explored unintentional changes in forces during performance of constant and cyclic force-production tasks (F-tasks) after visual feedback removal. Based on earlier studies, we expected all force parameters to drop exponentially with time. We also explored possible role of working memory in the force drop phenomena. Healthy subjects performed constant or cyclic isometric F-tasks with the index finger under visual feedback. The cyclic task was paced by a metronome. Removing visual feedback resulted in a consistent force drop in constant F-tasks and a qualitatively similar drift in the mean force in the cyclic F-task. Both were slow with characteristic times of about 10-20 s. In contrast, force amplitude in the cyclic F-task increased quickly (within 1-2 s). When the subjects were asked to stop producing force for 5 s after the visual feedback disappeared and then resume force production, no downward force drift was seen in constant F-tasks, while in cyclic F-tasks, the drift of the mean force was present and an exaggerated increase in force amplitude was also observed. We conclude that while working memory limitations may influence cyclic F-tasks, their role in determining the force drift in constant F-tasks is limited. The results of both experiments are interpreted within the referent configuration hypothesis supplemented with an idea of unintentional drift of referent coordinates (RC-back-coupling) induced by differences between the referent and actual body configurations.

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., (Published by Elsevier Ltd.)

We investigated the ability of two persons to produce force-stabilizing synergies in accurate multi-finger force production tasks under visual feedback on the total force only. The subjects produced a time profile of total force (the sum of two hand forces in one-person tasks and the sum of two subject forces in two-person tasks) consisting of a ramp-up, steady-state, and ramp-down segments; the steady-state segment was interrupted in the middle by a quick force pulse. Analyses of the structure of inter-trial finger force variance, motor equivalence, anticipatory synergy adjustments (ASAs), and the unintentional drift of the sharing pattern were performed. The two-person performance was characterized by a dramatically higher amount of inter-trial variance that did not affect total force, higher finger force deviations that did not affect total force (motor equivalent deviations), shorter ASAs, and larger drift of the sharing pattern. The rate of sharing pattern drift correlated with the initial disparity between the forces produced by the two persons (or two hands). The drift accelerated following the quick force pulse. Our observations show that sensory information on the task-specific performance variable is sufficient for the organization of performance-stabilizing synergies. They suggest, however, that two actors are less likely to follow a single optimization criterion as compared to a single performer. The presence of ASAs in the two-person condition might reflect fidgeting by one or both of the subjects. We discuss the characteristics of the drift in the sharing pattern as reflections of different characteristic times of motion within the subspaces that affect and do not affect salient performance variables.

We explored the phenomenon of unintentional changes in the apparent stiffness of the human arm produced by transient changes in the external force. The subjects performed a positional task against a constant baseline force and were instructed not to react to changes in the force. A HapticMaster robot produced a smooth force increase (a perturbation) leading to a hand movement, followed by a dwell time. No visible hand drift was observed during the dwell time. After the robot force dropped to its initial baseline value, the hand moved toward the initial position but stopped short of it. Small perturbations were applied at different time intervals along different directions during the dwell time. Arm apparent stiffness distribution in a horizontal plane was approximated with an ellipse. The apparent stiffness magnitude along the main axis of the ellipse showed a non-monotonic increase with dwell time, while the apparent stiffness along the minor axis did not change significantly. We interpreted the early part of the changes in the apparent stiffness as due to peripheral muscle properties. The later part is interpreted as caused by a combination of two processes: a drift in the referent hand coordinate due to the hypothesized back-coupling between the referent and actual hand coordinates and an implicit instruction to keep the hand steady when no changes in robot-generated force took place. The data provide support for the idea of back-coupling between the referent and actual body configurations, which may be an important contributor to stability of motor actions.

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., (Copyright © 2015 IBRO. Published by Elsevier Ltd. All rights reserved.)

Previous reports show that the forces produced by the fingers of one hand drop exponentially over time in the absence of visual feedback on the forces. We study the force production by the index fingers of both hands with no visual feedback. Subjects produced a specified total force with a specific contribution from each finger by pressing on force sensors. We observed that in the absence of visual feedback: (1) The finger forces dropped with time by an amount proportional to the magnitude of the initial force. For low initial force values (<7 % of MVC of individual finger force), the finger forces showed an increase; (2) the total force (sum of finger forces) evolution showed similar features; (3) finger forces changed in a way that facilitated more equitable force production by the two fingers; (4) all the force-time changes resemble exponential functions with similar time constants (~15 s). We propose that two processes interact to produce these patterns. (1) RC back-coupling: The central nervous system defines referent coordinates (RCs) for the digit tips, and the difference between the referent and actual coordinates leads to force production. If actual coordinates are not allowed to move to referent ones, referent coordinates show a slow drift toward the actual ones, leading to a force drop. (2) Sensory adaptation: This process, possibly related to sensory receptor characteristics, leads to an increase in finger force. RC back-coupling provides a common account of this and other reported phenomena of hand force or position changes across transient, external perturbations.

We explored stability of multi-finger cyclical accurate force production action by analysis of responses to small perturbations applied to one of the fingers and inter-cycle analysis of variance. Healthy subjects performed two versions of the cyclical task, with and without an explicit target. The "inverse piano" apparatus was used to lift/lower a finger by 1 cm over 0.5 s; the subjects were always instructed to perform the task as accurate as they could at all times. Deviations in the spaces of finger forces and modes (hypothetical commands to individual fingers) were quantified in directions that did not change total force (motor equivalent) and in directions that changed the total force (non-motor equivalent). Motor equivalent deviations started immediately with the perturbation and increased progressively with time. After a sequence of lifting-lowering perturbations leading to the initial conditions, motor equivalent deviations were dominating. These phenomena were less pronounced for analysis performed with respect to the total moment of force with respect to an axis parallel to the forearm/hand. Analysis of inter-cycle variance showed consistently higher variance in a subspace that did not change the total force as compared to the variance that affected total force. We interpret the results as reflections of task-specific stability of the redundant multi-finger system. Large motor equivalent deviations suggest that reactions of the neuromotor system to a perturbation involve large changes in neural commands that do not affect salient performance variables, even during actions with the purpose to correct those salient variables. Consistency of the analyses of motor equivalence and variance analysis provides additional support for the idea of task-specific stability ensured at a neural level.

The authors explored task-specific stability during accurate multifinger force production tasks with different numbers of instructed fingers. Subjects performed steady-state isometric force production tasks and were instructed not to interfere voluntarily with transient lifting-and-lowering perturbations applied to the index finger. The main results were (a) intertrial variance in the space of finger modes at steady states was larger within the subspace that had no effect on the total force (the uncontrolled manifold [UCM]); (b) perturbations caused large deviations of finger modes within the UCM (motor equivalence); and (c) deviations caused by the perturbation showed larger variance within the UCM. No significant effects of the number of task fingers were noted in any of the 3 indicators. The results are discussed within the frameworks of the UCM and referent configuration hypotheses. The authors conclude, in particular, that all the tasks were effectively 4-finger tasks with different involvement of task and nontask fingers.

The authors studied effects of practicing a 4-finger accurate force production task on multifinger coordination quantified within the uncontrolled manifold hypothesis. During practice, task instability was modified by changing visual feedback gain based on accuracy of performance. The authors also explored the retention of these effects, and their transfer to a prehensile task. Subjects practiced the force production task for 2 days. After the practice, total force variability decreased and performance became more accurate. In contrast, variance of finger forces showed a tendency to increase during the first practice session while in the space of finger modes (hypothetical commands to fingers) the increase was under the significance level. These effects were retained for 2 weeks. No transfer of these effects to the prehensile task was seen, suggesting high specificity of coordination changes. The retention of practice effects without transfer to a different task suggests that further studies on a more practical method of improving coordination are needed.

We investigated stability of action by a multifinger system with three methods: analysis of intertrial variance, application of transient perturbations, and analysis of the system's motion in different state spaces. The "inverse piano" device was used to apply transient (lifting-and-lowering) perturbations to individual fingers during single- and two-finger accurate force production tasks. In each trial, the perturbation was applied either to a finger explicitly involved in the task or one that was not. We hypothesized that, in one-finger tasks, task-specific stability would be observed in the redundant space of finger forces but not in the nonredundant space of finger modes (commands to explicitly involved fingers). In two-finger tasks, we expected that perturbations applied to a nontask finger would not contribute to task-specific stability in mode space. In contrast to our expectations, analyses in both force and mode spaces showed lower stability in directions that did not change total force output compared with directions that did cause changes in total force. In addition, the transient perturbations led to a significant increase in the enslaving index. We consider these results within a theoretical scheme of control with referent body configurations organized hierarchically, using multiple few-to-many mappings organized in a synergic way. The observed volatility of enslaving, greater equifinality of total force compared with elemental variables, and large magnitude of motor equivalent motion in both force and mode spaces provide support for the concept of task-specific stability of performance and the existence of multiple neural loops, which ensure this stability., (Copyright © 2014 the American Physiological Society.)

The effects of muscle fatigue on the stability of precision grasps are not well known. The purpose of the current study was to investigate the effects of exercise-induced fatigue of a digit on prehension synergies in a static precision grasp. One group of participants performed the fatiguing exercise using the thumb (group-thumb) and the second group performed the exercise using the index finger (group-index). Grasp force and load-resisting force-stabilizing synergies were weaker during fatigue for group-thumb and showed no significant change for group-index. These results indicate that fatiguing the thumb compromises the stability of the precision grasp more than when the index finger is fatigued. Our results support the idea of hierarchical organization of prehension control. We proffer an explanation of our results based on two control constructs: a) Principle of superposition. This principle states that prehension can be viewed as a superposition of two independent processes controlling the slip and the tilt of the object respectively; and b) The referent configuration hypothesis. According to this hypothesis, the neural control of actions is associated with defining a set of referent values for task-related coordinates (given an external force field) defined as the referent configuration.

In this study, we explored relations between indices of postural sway and perceived comfort during pointing postures performed by standing participants. The participants stood on a force plate, grasped a pointer with the dominant (right) hand, and pointed to targets located at four positions and at two distances from the body. We quantified postural sway over 60-s intervals at each pointing posture, and found no effects of target location or distance on postural sway indices. In contrast, comfort ratings correlated significantly with indices of one of the sway components, trembling. Our observations support the hypothesis that rambling and trembling sway components involve different neurophysiological mechanisms. They also suggest that subjective perception of comfort may be more important than the actual posture for postural sway., (Copyright © 2014 Elsevier Ireland Ltd. All rights reserved.)

The extrinsic digit muscles naturally couple wrist action and grip force in prehensile tasks. We explored the effects of wrist position on the steady-state grip force and grip-force change during imposed changes in the grip aperture [apparent stiffness (AS)]. Subjects held an instrumented handle steady using a prismatic five-digit grip. The grip aperture was changed slowly, while the subjects were instructed not to react voluntarily to these changes. An increase in the aperture resulted in an increase in grip force, and its contraction resulted in a proportional drop in grip force. The AS values (between 4 and 6 N/cm) were consistent across a wide range of wrist positions. These values were larger when the subjects performed the task with eyes open as compared to eyes-closed trials. They were also larger for trials that started from a larger initial aperture. After a sequence of aperture increase and decrease to the initial width, grip force dropped by about 25% without the subjects being aware of this. We interpret the findings within the referent configuration hypothesis of grip-force production. The results support the idea of back-coupling between the referent and actual digit coordinates. According to this idea, the central nervous system defines referent coordinates for the digit tips, and the difference between the referent and actual coordinates leads to force production. If actual coordinates are not allowed to move to referent ones, referent coordinates show a relatively slow drift toward the actual ones.

This study was motivated by the double action of extrinsic hand muscles that produce grip force and also contribute to wrist torque. We explored interactions between grip force and wrist torque in isometric force production tasks. In particular, we tested a hypothesis that an intentional change in one of the two kinetic variables would produce an unintentional change in the other (enslaving). When young healthy subjects produced accurate changes in the grip force, only minor effects on the force produced by the hand (by wrist flexion/extension action) were observed. In contrast, a change in the hand force produced consistent changes in grip force in the same direction. The magnitude of such unintentional grip force change was stronger for intentional hand force decrease as compared to hand force increase. These effects increased with the magnitude of the initial grip force. When the subjects were asked to produce accurate total force computed as the sum of the hand and grip forces, strong negative covariation between the two forces was seen across trials interpreted as a synergy stabilizing the total force. An index of this synergy was higher in the space of "modes," hypothetical signals to the two effectors that could be changed by the controller one at a time. We interpret the complex enslaving effects (positive force covariation) as conditioned by typical everyday tasks. The presence of synergic effects (negative, task-specific force covariation) can be naturally interpreted within the referent configuration hypothesis.

During single-finger force production, the non-instructed fingers unintentionally produce force (finger enslaving). In this study, enslaving effects were compared between the dominant and non-dominant hands. The test consisted of a series of maximum voluntary contractions with different finger combinations. Enslaving matrices were calculated by means of training an artificial neural network. The dominant hand was found to be stronger, but there was found to be no difference between the overall enslaving effects in the dominant and non-dominant hands. There was no correlation between the magnitude of finger enslaving and the performance in such tests as the Edinburgh Handedness Inventory, the Grooved Pegboard test, and the Jebsen-Taylor Hand Function test. Each one of those three tests showed a significant difference between the dominant and non-dominant hand performances. Eleven subjects were retested after two months, and it was found that enslaving effects did not fluctuate significantly between the two testing sessions. While the dominant and non-dominant hands are involved differently in everyday tasks, e.g. in writing or eating, this practice does not cause significant differences in enslaving between the hands., (Copyright © 2013 Elsevier B.V. All rights reserved.)

Background: Mechanical properties of human digits may have significant implications for the hand function. We quantified several mechanical characteristics of individual digits in young and older adults., Methods: Digit tip friction was measured at several normal force values using a method of induced relative motion between the digit tip and the object surface. A modified quick-release paradigm was used to estimate digit apparent stiffness, damping, and inertial parameters. The subjects grasped a vertical handle instrumented with force/moment sensors using a prismatic grasp with four digits; the handle was fixed to the table. Unexpectedly, one of the sensors yielded leading to a quick displacement of the corresponding digit. A second-order, linear model was used to fit the force/displacement data., Findings: Friction of the digit pads was significantly lower in older adults. The apparent stiffness coefficient values were higher while the damping coefficients were lower in older adults leading to lower damping ratio. The damping ratio was above unity for most data in young adults and below unity for older adults. Quick release of a digit led to force changes in other digits of the hand, likely due to inertial hand properties. These phenomena of "mechanical enslaving" were smaller in older adults although no significant difference was found in the inertial parameter in the two groups., Interpretations: The decreased friction and damping ratio present challenges for the control of everyday prehensile tasks. They may lead to excessive digit forces and low stability of the grasped object., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The authors studied effects of healthy aging on 3 components of the internal force vector during static prehensile tasks. Young and older subjects held an instrumented handle using a 5-digit prismatic grasp under different digit configurations and external torques. Across digit configurations, older subjects showed larger internal normal (grip) and tangential (load-resisting) digit force components and larger internal moment of force. In contrast to earlier reports, safety margin values were not higher in the older subjects. The results show that the increased grip force in older persons is a specific example of a more general age-related problem reflected in the generation of large internal force vectors in prehensile tasks. It is possible that the higher internal forces increase the apparent stiffness of the hand+handle system and, hence, contribute to its stability. This strategy, however, may be maladaptive, energetically wasteful, and inefficient in ensuring safety of hand-held objects.

A hypothesis was proposed that the central nervous system controls force production by the fingers through hypothetical neural commands. The neural commands are scaled between values of 0 to 1, indicating no intentional force production or maximal voluntary contraction (MVC) force production, respectively. A matrix of interfinger connections transforms neural commands into finger forces. Two methods have been proposed to compute the interfinger connection matrix. The first method uses only single finger MVC trials and multiplies the interfinger connection matrix by a gain factor. The second method uses a neural network model based on experimental data. The performance of the two methods was compared on the MVC data and on a data set of submaximal forces, collected over a range of total forces and moments of force. The methods were compared in terms of (1) ability to predict finger forces, (2) accuracy of neural command reconstruction, and (3) preserved planarity of force data for submaximal force production task. Both methods did a reasonable job of predicting the total force in multifinger MVC trials; however, the neural network model performed better in regards to all other criteria. Overall, the results indicate that for modeling multifinger interaction the neural network method is preferable.

During power grasp, the number of local force maxima reflects either the central nervous system's preferential use of particular hand regions, or anatomical constraints, or both. Previously, both bimodal and trimodal force maxima have been hypothesized for power grasp of a cylindrical handle. Here we measure the number of local force maxima, with a resolution of 4.8°, when performing pushing and pulling efforts in the plane perpendicular to the cylinder's long axis. Twelve participants produced external forces to eight targets. The number of contacts was defined as the number of local maxima exceeding background variance. A minimum of four and a maximum of five discrete contacts were observed in all subjects at the distal phalanges and metacarpal heads. We thus reject previous hypotheses of bimodal or trimodal force control for cylindrical power grasping. Since we presently observed only 4-5 contacts, which is rather low considering the hand's kinematic flexibility in the flexion plane, we also reject hypotheses of continuous contact, which are inherent to current grasping taxonomy. A modification to current grasping taxonomy is proposed wherein power grasp contains separate branches for continuous and discrete contacts, and where power and precision grasps are distinguished only by grasp manipulability.

We explored a hypothesis that transient perturbations applied to a redundant system result in equifinality in the space of task-related performance variables but not in the space of elemental variables. The subjects pressed with four fingers and produced an accurate constant total force level. The "inverse piano" device was used to lift and lower one of the fingers smoothly. The subjects were instructed "not to intervene voluntarily" with possible force changes. Analysis was performed in spaces of finger forces and finger modes (hypothetical neural commands to fingers) as elemental variables. Lifting a finger led to an increase in its force and a decrease in the forces of the other three fingers; the total force increased. Lowering the finger back led to a drop in the force of the perturbed finger. At the final state, the sum of the variances of finger forces/modes computed across repetitive trials was significantly higher than the variance of the total force/mode. Most variance of the individual finger force/mode changes between the preperturbation and postperturbation states was compatible with constant total force. We conclude that a transient perturbation applied to a redundant system leads to relatively small variance in the task-related performance variable (equifinality), whereas in the space of elemental variables much more variance occurs that does not lead to total force changes. We interpret the results within a general theoretical scheme that incorporates the ideas of hierarchically organized control, control with referent configurations, synergic control, and the uncontrolled manifold hypothesis.

We investigated the effects of fatigue produced by timed maximal voluntary contraction (MVC) of the index finger of the right hand on performance in MVC and accurate cyclic force production tasks in right-handed young (Young group) and strength-matched elderly (Elderly group) participants. We hypothesized that, before fatigue, the Elderly group would show weaker force-stabilizing synergies and smaller adaptive changes in the synergy index during fatigue. Synergies were defined as covaried adjustments of neural commands to fingers (finger modes) across trials that stabilize total force. Fatigue caused a significant reduction in the MVC, which was larger in the Young group compared with the Elderly group for both fatigued finger (index finger) and four fingers (index, middle, ring, and little fingers pressing together). Indexes of finger enslaving (lack of individuation) increased with fatigue in both groups. The index of force-stabilizing synergies was similar for the two groups before fatigue, while its increase with fatigue was significantly larger in the Elderly group compared with the Young group. We infer that changes in the indexes of finger interaction (enslaving) and coordination (synergy) with age seem to be correlated with changes in muscle strength. This correlation may be causally related to the progressive death of neurons at different levels of the neuromotor hierarchy. The surprisingly large changes in the synergy index with fatigue in older adults suggest that, by itself, aging does not necessarily lead to impairment in synergic control. Strength training may be a method to avoid age-related decrement in 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.

Extrinsic digit muscles contribute to both fingertip forces and wrist movements (FDP and FPL-flexion, EDC-extension). Hence, it is expected that finger forces depend on the wrist movement and position. We investigated the relation between grip force and wrist kinematics to examine whether and how the force (1) scales with wrist flexion-extension (FE) angle and (2) can be predicted from accelerations induced during FE movement. In one experiment, subjects naturally held an instrumented handle using a prismatic grasp and performed very slow FE movements. In another experiment, the same movement was performed cyclically at three prescribed frequencies. In quasistatic conditions, the grip force remained constant over the majority of the wrist range of motion. During the cyclic movements, the grip force changed. The changes were described with a linear regression model that represents the thumb and virtual finger (VF = four fingers combined) normal forces as the sum of the effects of the object's tangential and radial accelerations and an object-weight-dependent constant term. The model explained 99 % of the variability in the data. The independence of the grip force from wrist position agrees with the theory that the thumb and VF forces are controlled with two neural variables that encode referent coordinates for each digit while accounting for changes in the position dependence of muscle forces, rather than a single neural variable like referent aperture. The results of the cyclical movement study extend the principle of superposition (some complex actions can be decomposed into independently controlled elemental actions) for a motor task involving simultaneous grip-force exertion and wrist motion with significant length changes of the grip-force-producing muscles.

We studied a mechanism of feed-forward control of a multi-finger action, namely anticipatory synergy adjustments (ASAs), prior to a quick force correction in response to a change in the gain of the visual feedback. Synergies were defined as co-varied across trials adjustments of commands to fingers that stabilized (decreased variance of) the total force. We hypothesized that ASAs would be highly sensitive to prior information about the timing of the action but not to information on its direction, i.e., on whether the gain would go up or down. The subjects produced accurate constant total force by pressing with four fingers on individual force sensors. The feedback signal could change from veridical (the sum of finger forces) to modified, with the middle finger force multiplied by 0.2 or by 1.8. The timing of the gain change and its direction could be known or unknown to the subject in advance. When the timing of the gain change was known, ASA was seen as a drop in the synergy index starting about 250-300 ms prior to the first visible correction of the total force. When the gain change timing was unknown, ASAs started much later, less than 100 ms prior to the total force correction. The magnitude of synergy index changes was significantly larger under the "time known" conditions. Information on the direction of the visual gain change had no effect on the ASA timing, while the ASA magnitude was somewhat larger when this information was not available to the subject. After the total force correction, the synergy index was significantly larger for the force signal computed using the modified gain values as compared to the synergy index value for the actual total force. We conclude that ASAs represent an important feed-forward motor control mechanism that allows preparing for a quick action even when the direction of the action is not known in advance. The results emphasize the subtle control of multi-finger synergies that are specific to the exact contributions of individual fingers to performance variables. The data fit well the central back-coupling hypothesis of synergies and the idea of control with referent body configurations.

We investigated the effects of exercise-induced fatigue of a digit on the biomechanics of a static prehension task. The participants were divided into two groups. One group performed the fatiguing exercise using the thumb (group-thumb) and the second group performed the exercise using the index finger (group-index). We analyzed the prehensile action as being based on a two-level hierarchy. Our first hypothesis was that fatigue of the thumb would have stronger effects at the upper level (action shared between the thumb and all four fingers combined-virtual finger) and fatigue of the index finger would have stronger effects at the lower level of the hierarchy (action of the virtual finger shared among actual fingers). We also hypothesized that fatigue would cause a decrease in the normal force applied by the exercised digit and correspondingly lead to a decrease in the normal force applied by the opposing digit(s). Our third hypothesis was that fatigue would leave the tangential forces unaffected. Fatigue led to a significant drop in the normal force of both exercised and non-exercised (opposing) digits. The tangential forces of the exercised digits increased after fatigue. This led to a drop in the safety margin in the group-thumb, but not group-index. As such, the results supported the first two hypotheses but not the third hypothesis. Overall, the results suggested that fatigue triggered a chain reaction that involved both forces and moments of force produced by individual digits leading to a violation of the principle of superposition. The findings are interpreted within the framework of the referent configuration hypothesis.

We studied the effects of a single practice session of a variable task with subject-specific adjustments of task difficulty (instability) on indices of multi-finger coordination in young and elderly persons. The main hypothesis was that practicing such a task would lead to contrasting changes in the amounts of two components of variance estimated across repetitive trials within the uncontrolled manifold (UCM) hypothesis: V UCM that had no effect on total force and V ORT that affected total force. In addition, we also expected to see strong transfer effects to a different task. A variable task with graded instability was designed to encourage use of variable solutions during the accurate production of total force with two fingers. The subjects practiced with the index and middle fingers pressing on individual force sensors. Overall, the older subjects showed lower indices of performance and higher indices of both V UCM and V ORT. After about 1 h of practice, both groups showed an increase in the index of involuntary force production by non-task fingers (enslaving). Both groups improved the indices of performance. The two variance indices showed opposite effects of practice: V ORT dropped with practice, while V UCM increased leading to an increase in the total amount of variance in the space of commands to fingers and in the index of force-stabilizing synergy. Performance in a simpler, non-practiced task improved, but there was no transfer of the changes in the structure of variance. Specifically, both variance components, V ORT and V UCM, dropped in the non-practiced task. The results show that the neural system responsible for synergies stabilizing important features of performance is highly adaptable to practice of tasks designed to encourage use of variable solutions. We view the results as highly promising for future use in populations with impaired coordination characterized by low synergy indices.

This study joined two approaches to motor control. The first approach comes from cognitive psychology and is based on the idea that goal postures and movements are chosen to satisfy task-specific constraints. The second approach comes from the principle of motor abundance and is based on the idea that control of apparently redundant systems is associated with the creation of multi-element synergies stabilizing important performance variables. The first approach has been tested by relying on psychophysical ratings of comfort. The second approach has been tested by estimating variance along different directions in the space of elemental variables such as joint postures. The two approaches were joined here. Standing subjects performed series of movements in which they brought a hand-held pointer to each of four targets oriented within a frontal plane, close to or far from the body. The subjects were asked to rate the comfort of the final postures, and the variance of their joint configurations during the steady state following pointing was quantified with respect to pointer endpoint position and pointer orientation. The subjects showed consistent patterns of comfort ratings among the targets, and all movements were characterized by multi-joint synergies stabilizing both pointer endpoint position and orientation. Contrary to what was expected, less comfortable postures had higher joint configuration variance than did more comfortable postures without major changes in the synergy indices. Multi-joint synergies stabilized the pointer position and orientation similarly across a range of comfortable/uncomfortable postures. The results are interpreted in terms conducive to the two theoretical frameworks underlying this work, one focusing on comfort ratings reflecting mean postures adopted for different targets and the other focusing on indices of joint configuration variance.

Background: Carpal tunnel syndrome is a disorder caused by increased pressure in the carpal tunnel associated with repetitive, stereotypical finger actions. Little is known about in vivo geometrical changes in the carpal tunnel caused by motion at the finger joints and exerting a fingertip force., Methods: The hands and forearms of five subjects were scanned using a 3.0 T magnetic resonance imaging scanner. The metacarpophalangeal joint of the index finger was placed in: flexion, neutral and extension. For each joint posture subjects either produced no active force (passive condition) or exerted a flexion force to resist a load (~4.0 N) at the fingertip (active condition). Changes in the radii of curvature, position and transverse plane area of the flexor digitorum profundus tendons at the carpal tunnel level were measured., Results: The radius of curvature of the flexor digitorum profundus tendons, at the carpal tunnel level, was significantly affected by posture of the index finger metacarpophalangeal joint (P<0.05) and the radii was significantly different between fingers (P<0.05). Actively producing force caused a significant shift (P<0.05) in the flexor digitorum profundus tendons in the ventral (palmar) direction. No significant change in the area of an ellipse containing the flexor digitorum profundus tendons was observed between conditions., Interpretation: The results show that relatively small changes in the posture and force production of a single finger can lead to significant changes in the geometry of all the flexor digitorum profundus tendons in the carpal tunnel. Additionally, voluntary force production at the fingertip increases the moment arm of the FDP tendons about the wrist joint., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The central hypothesis explored in the experiment is that adjustments of fingertip force vectors during object manipulation could result from a simple scaling rule applied to commands to individual digits. The commands have been associated with referent coordinates of the digit tips. The subjects performed quick lifting movements (over 20 cm in under 0.5 s) of a horizontally oriented handle with different combinations of the external load and torque. The prismatic grasp was used with the 4 fingers pressing on the bottom of the handle and the thumb acting on its top. Principal component and correlation analyses applied to the normal and tangential force vector components confirmed that the force direction of each digit was kept nearly constant in the object-centered referent frame across the loading conditions and movement phases. The middle and ring fingers showed weaker correlations between the force components as compared to the index and little fingers. The differences were likely related to the different roles of the normal force components in the moment of force production. The neural control of the hand, within the studied task, may be adequately described as a simple rule applied to a handful of parameters, such as the referent digit-tip coordinates.

The neural control of movement has been described using different sets of elemental variables. Two possible sets of elemental variables have been suggested for finger pressing tasks: the forces of individual fingers and the finger commands (also called finger modes or central commands). The authors analyzed which of the 2 sets of the elemental variables is more likely used in the optimization of the finger force sharing and which set is used for the stabilization of performance. They used two recently developed techniques-the analytical inverse optimization (ANIO) and the uncontrolled manifold (UCM) analysis-to evaluate each set of elemental variables with respect to both aspects of performance. The results of the UCM analysis favored the finger commands as the elemental variables used for performance stabilization, while ANIO worked equally well on both sets of elemental variables. A simple scheme is suggested as to how the CNS could optimize a cost function dependent on the finger forces, but for the sake of facilitation of the feed forward control it substitutes the original cost function by a cost function, which is convenient to optimize in the space of finger commands.

We investigated the effect of fatigue produced by timed maximal voluntary contraction (MVC) of the index finger of one of the hands on performance in MVC and accurate cyclic force production tasks in right-handed subjects. Based on earlier studies, we hypothesized that fatigue would produce an increase in the indices of force-stabilizing synergies in both hands as well as between the hands in two-hand tasks. Synergies were defined as co-varied adjustments of commands to fingers (modes) across cycles that stabilized total force. Fatigue caused a significant reduction in the MVC of the exercised as well as the non-exercised hand. Indices of finger enslaving (lack of individuation) increased with fatigue in both hands, although the increase was significant in the exercised hand only. In contrast to the significant effects of fatigue on MVC forces performed by the non-exercised hand, there were no comparable transfer effects on the root mean square errors during accurate force production. During one-hand tasks, both hands showed high indices of force-stabilizing synergies. These indices were larger in the left hand. Fatigue led to a general increase in synergy indices. Exercise by the left hand had stronger effects on synergy indices seen in both hands. Exercise by the right hand showed ipsilateral effects only. Smaller effects of fatigue were observed on accuracy of performance of the force-down segments of the force cycles compared to the force-up segments. For the bimanual tasks, synergies were analyzed at two hierarchical levels, two-hand (four-finger) and within-a-hand (two-finger). An increase in the synergy index with fatigue was observed at the lower (two-finger) level of the hierarchy only. We interpret the lack of effects of fatigue at the upper (two-hand) level as a consequence of a trade-off between synergies at different levels of the hierarchy. The differences between the hands are discussed within the dynamic dominance hypothesis., (Copyright © 2012 Elsevier B.V. All rights reserved.)

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.

We explored digit coordination during the acceleration phase of a quick lifting movement of a hand-held horizontal object. We tested three hypotheses related to: (1) the scaling of mechanical variables produced by the hand with changes in the external load, torque, and moment of inertia; (2) changes in the safety margin for the thumb with both the loading conditions and acceleration; and (3) changes in the indices of synergies. The subjects held a horizontal handle with a prismatic grasp (the thumb acted on top of the handle) and performed series of "very quick" lifting movements to a visual target. Multi-digit synergies were quantified as co-variation indices among elemental variables (forces and moments produced by individual digits). The resultant force scaled with the external load but not torque, while the grip force scaled with the external torque but not load. The safety margin dropped with an increase in acceleration; it also showed changes with the external torque and moment of inertia. Total moment of force was primarily produced by the tangential forces (over 80 %) across all movement phases and loading conditions. The index and little fingers produced close to zero moment with their normal forces, while the middle and ring fingers produced consistent moments due to the reproducible shifts of their centers of pressure. Synergy indices at the upper level of the assumed hierarchy (the task is shared between the thumb and virtual finger--an imagined digit with the action equal to that of the four fingers combined) did not drop with acceleration for the three force vector components and one of the moment vector components. They did drop with acceleration at the lower level (virtual finger action is shared among the four fingers). There was a trade-off between synergy indices computed at the two levels for the three force vector components, but not for the moment of force components. We confirmed specialization of different fingers with respect to different task components in quick manipulation tasks. The findings have implications for hypotheses on the control of voluntary movements involving redundant sets of effectors. Within the referent configuration hypothesis, components of a referent configuration may be adjusted to task mechanical characteristics using simple scaling rules. The neural organization of multi-digit synergies in a hierarchal system is able to selectively protect synergies related to stabilization of some performance variables from detrimental effects of the rate of change of those variables. A large number of apparently redundant elemental variables are not the source of additional computational problems but may be beneficial, allowing the central nervous system to facilitate synergies at both levels of the hierarchy.

Background: The purpose of this study was to use magnetic resonance imaging to measure the moment arm of the flexor digitorum superficialis tendon about the metacarpophalangeal joint of the index, middle, ring, and little fingers when the position and force production level of the index finger was altered. A secondary goal was to create regression models using anthropometric data to predict moment arms of the flexor digitorum superficialis about the metacarpophalangeal joint of each finger., Methods: The hands of subjects were scanned using a 3.0 T magnetic resonance imaging scanner. The metacarpophalangeal joint of the index finger was placed in: flexion, neutral, and extension. For each joint configuration subjects produced no active force (passive condition) and exerted a flexion force to resist a load at the fingertip (active condition)., Results: The following was found: (1) The moment arm of the flexor digitorum superficialis at the metacarpophalangeal joint of the index finger (a) increased with the joint flexion and stayed unchanged with finger extension; and (b) decreased with the increase of force at the neutral and extended finger postures and did not change at the flexed posture. (2) The moment arms of the flexor digitorum superficialis tendon of the middle, ring, and little fingers (a) did not change when the index metacarpophalangeal joint position changed (P>0.20); and (b) The moment arms of the middle and little fingers increased when the index finger actively produced force at the flexed metacarpophalangeal joint posture. (4) The moment arms showed a high correlation with anthropometric measurements., Interpretation: Moment arms of the flexor digitorum superficialis change due to both changes in joint angle and muscle activation; they scale with various anthropometric measures., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

We investigated age-related differences in finger coordination during rotational hand actions. Two hypotheses based on earlier studies were tested: higher safety margins and lower synergy indices were expected in the elderly. Young and elderly subjects held a handle instrumented with five six-component force sensors and performed discrete accurate pronation and supination movements. The weight of the system was counterbalanced with another load. Indices of synergies stabilizing salient performance variables, such as total normal force, total tangential force, moments produced by these forces, and total moment of force were computed at two levels of a hypothetical control hierarchy, at the virtual finger-thumb level and at the individual finger level. At each level, synergy indices reflected the normalized difference between the sum of the variances of elemental variables and variance of their combined output, both computed at comparable phases over repetitive trials. The elderly group performed the task slower and showed lower safety margins for the thumb during the rotation phase. Overall, the synergy indices were not lower in the elderly group. In several cases, these indices were significantly higher in the elderly than in the younger participants. Hence, both main hypotheses have been falsified. We interpret the unexpectedly low safety margins in the elderly as resulting from several factors such as increased force variability, impaired feed-forward control, and the fact that there was no danger of dropping the object. Our results suggest that in some natural tasks, such as the one used in this study, healthy elderly persons show no impairment, as compared to younger persons, in their ability to organize digits into synergies stabilizing salient performance variables., (Copyright © 2011 Elsevier B.V. All rights reserved.)

When grasping and manipulating objects, the central controller utilizes the mechanical advantage of the normal forces of the fingers for torque production. Whether the same is valid for tangential forces is unknown. The main purpose of this study was to determine the patterns of finger tangential forces and the use of mechanical advantage as a control mechanism when dealing with objects of nonuniform finger positioning. A complementary goal was to explore the interaction of mechanical advantage (moment arm) and the role a finger has as a torque agonist/antagonist with respect to external torques (±0.4 N m). Five 6-df force/torque transducers measured finger forces while subjects held a prism handle (6 cm width × 9 cm height) with and without a single finger displaced 2 cm (handle width). The effect of increasing the tangential moment arm was significant (p < .01) for increasing tangential forces (in >70% of trials) and hence creating greater moments. Thus, the data provides evidence that the grasping system as a rule utilizes mechanical advantage for generating tangential forces. The increase in tangential force was independent of whether the finger was acting as a torque agonist or antagonist, revealing their effects to be additive.

We used two methods to address two aspects of multi-finger synergies and their changes after fatigue of the index finger. Analytical inverse optimization (ANIO) was used to identify cost functions and corresponding spaces of optimal solutions over a broad range of task parameters. Analysis within the uncontrolled manifold (UCM) hypothesis was used to quantify co-variation of finger forces across repetitive trials that helped reduce variability of (stabilized) performance variables produced by all the fingers together. Subjects produced steady-state levels of total force and moment of force simultaneously as accurately as possible by pressing with the four fingers of the right hand. Both before and during fatigue, the subjects performed single trials for many force-moment combinations covering a broad range; the data were used for the ANIO analysis. Multiple trials were performed at two force-moment combinations; these data were used for analysis within the UCM hypothesis. Fatigue was induced by 1-min maximal voluntary contraction exercise by the index finger. Principal component (PC) analysis showed that the first two PCs explained over 90% of the total variance both before and during fatigue. Hence, experimental observations formed a plane in the four-dimensional finger force space both before and during fatigue conditions. Based on this finding, quadratic cost functions with linear terms were estimated from the experimental data. The dihedral angle between the plane of optimal solutions and the plane of experimental observations (D (ANGLE)) was very small (a few degrees); it increased during fatigue. There was an increase in fatigue of the coefficient at the quadratic term for the index finger force balanced by a drop in the coefficients for the ring and middle fingers. Within each finger pair (index-middle and ring-little), the contribution of the "central" fingers to moment production increased during fatigue. An index of antagonist moment production dropped with fatigue. Fatigue led to higher co-variation indices during pronation tasks (index finger is an agonist) but opposite effects during supination tasks. The results suggest that adaptive changes in co-variation indices that help stabilize performance may depend on the role of the fatigued element, agonist or antagonist.

Radial force (F(r)) distributions describe grip force coordination about a cylindrical object. Recent studies have employed only explicit F(r) tasks, and have not normalized for anatomical variance when considering F(r) distributions. The goals of the present study were (i) to explore F(r) during tangential force production tasks, and (ii) to examine the extent to which anatomical registration (i.e. spatial normalization of anatomically analogous structures) could improve signal detectability in F(r) data. Twelve subjects grasped a vertically oriented cylindrical handle (diameter=6 cm) and matched target upward tangential forces of 10, 20, and 30 N. F(r) data were measured using a flexible pressure mat with an angular resolution of 4.8°, and were registered using piecewise-linear interpolation between five manually identified points-of-interest. Results indicate that F(r) was primarily limited to three contact regions: the distal thumb, the distal fingers, and the fingers' metatacarpal heads, and that, while increases in tangential force caused significant increases in F(r) for these regions, they did not significantly affect the F(r) distribution across the hand. Registration was found to substantially reduce between-subject variability, as indicated by both accentuated F(r) trends, and amplification of the test statistic. These results imply that, while subjects focus F(r) primarily on three anatomical regions during cylindrical grasp, inter-subject anatomical differences introduce a variability that, if not corrected for via registration, may compromise one's ability to draw anatomically relevant conclusions from grasping force data., (Copyright © 2011 Elsevier Ltd. All rights reserved.)