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

Reschechtko S, Zatsiorsky VM, Latash ML. Stability of multifinger action in different state spaces. J Neurophysiol 112: 3209-3218, 2014. First published September 24, 2014 doi:10.1152/jn.00395.2014.-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-andlowering) perturbations to individual fingers during single-and twofinger 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. redundancy; abundance; synergy; referent configuration; uncontrolled manifold hypothesis; equifinality; enslaving ONE OF THE FUNDAMENTAL PROBLEMS of motor control is that of redundancy (Bernstein 1967): at any level of description of the system involved in movement production, the number of elements (and of their output variables, elemental variables) is higher than the number of constraints associated with typical motor tasks. Recently, the problem of motor redundancy has been revisited in the context of the principle of motor abundance (Gelfand and Latash 1998;Latash 2012). This approach posits a way for the central nervous system (CNS) to take advantage of all of the elemental variables at its disposal to ensure stability of movement, rather than requiring the CNS to achieve a single, optimal solution by eliminating redundant degrees of freedom. Thus the principle of abundance avoids requiring computational, and physiologically unfeasible, control schemes, which involve explicit instruction to all body structures involved in a given action.A theoretical scheme has been developed, which unites the principle of abundance with the idea of hierarchical...

Background After a single slip, older adults rapidly make adaptive changes to avoid or eliminate further backward loss of balance or a fall. This rapid adaptation has been termed the “single trial effect”. The purpose of this study was to explore the relationship between the motor errors subjects experienced upon a novel slip and the selection and execution of corrective response by which they modified their ongoing gait pattern and turned it into a protective step. Methods A forward slip was induced in the laboratory among 145 community-living older (≥65 year old) adults who were protected by an overhead full body harness system. An eight-camera motion analysis system recorded subjects’ kinematics, which was used to compute their instability (motor error), recovery step placement (response selection), and stability gain (motor correction). Findings A linear relationship was found between the stability errors at recovery foot liftoff and the distance between the recovery foot and slipping foot at the time of its touchdown, reflecting an appropriate selection of response that was proportionate to the motor error. A linear relationship was also found between this step modification and resulting stability gain, indicating that greater step modification resulted in greater stability gain. This learning behavior was surprisingly consistent regardless whether the outcome was a recovery or a fall. Interpretations These results suggest that fallers and non-fallers all have an intact motor learning foundation that has enabled them to rapidly improve their stability in subsequent exposures.

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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 sub-spaces that affect and do not affect salient performance variables.

We investigated multi-digit 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 (UCM) framework by quantifying the inter-trial 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 inter-trial 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.

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.

The study explored unintentional force changes elicited by removing visual feedback during cyclical, two-finger isometric force production tasks. Subjects performed two types of tasks at 1 Hz, paced by an auditory metronome. One – Force task – required cyclical changes in total force while maintaining the sharing, defined as relative contribution of a finger to total force. The other task – Share task – required cyclical changes in sharing while keeping total force unchanged. Each trial started under full visual feedback on both force and sharing; subsequently, feedback on the variable that was instructed to stay constant was frozen, and finally feedback on the other variable was also removed. In both tasks, turning off visual feedback on total force elicited a drop in the mid-point of the force cycle and an increase in the peak-to-peak force amplitude. Turning off visual feedback on sharing led to a drift of mean share toward 50:50 across both tasks. Without visual feedback there was consistent deviation of the two force time series from the in-phase pattern (typical of the Force task) and from the out-of-phase pattern (typical of the Share task). This finding is in contrast to most earlier studies that demonstrated only two stable patterns, in-phase and out-of-phase. We interpret the results as consequences of drifts of parameters in a dynamical system leading in particular to drifts in the referent finger coordinates toward their actual coordinates. The relative phase desynchronization is caused by the right-left differences in the hypothesized drift processes, consistent with the dynamic dominance hypothesis.