Reach Corrections Toward Moving Objects are Faster Than Reach Corrections Toward Instantaneously Switching Targets.

Visually guided reaching is a common motor behavior that engages subcortical circuits to mediate rapid corrections. Although these neural mechanisms have evolved for interacting with the physical world, they are often studied in the context of reaching toward virtual targets on a screen. These targets often change position by disappearing from one place reappearing in another instantaneously. In this study, we instructed participants to perform rapid reaches to physical objects that changed position in different ways. In one condition, the objects moved very rapidly from one place to another. In the other condition, illuminated targets instantaneously switched position by being extinguished in one position and illuminating in another. Participants were consistently faster in correcting their reach trajectories when the object moved continuously.

Postural sway is not affected by estrogen fluctuations during the menstrual cycle.

When people stand still, they exhibit a phenomenon called postural sway, or spontaneous movement of the body's center of pressure, which is related to balance control. In general females show less sway than males, but this difference only begins to appear around puberty, pointing to different levels of sex hormones as one potential mechanism for sway sex differences. In this study, we followed cohorts of young females using oral contraceptives (n = 32) and not using oral contraceptives (n = 19), to investigate associations between estrogen availability and postural sway. All participants visited the lab four times over the putative 28-day menstrual cycle. At each visit, we performed blood draws to measure plasma estrogen (estradiol) levels, and tests of postural sway using a force plate. During late follicular and mid-luteal phase, estradiol levels were lower in participants using oral contraceptives (mean differences [95% CI], respectively: -231.33; [-800.44, 337.87]; -613.26; [-1333.60, 107.07] pmol/L; main effect p < 0.001), reflecting expected consequences of oral contraceptive use. Despite these differences, postural sway was not significantly different between participants who were using oral contraceptives and participants who were not (mean difference: 2.09 cm; 95% CI = [-1.05, 5.22]; p = 0.132). Overall, we found no significant effects of the estimated menstrual cycle phase-or absolute levels of estradiol-on postural sway.

Stretch reflexes.

Many of us know about stretch reflexes from the doctor's office, when a physician taps the tendon near our kneecap to elicit a quick knee extension. This procedure is used as a diagnostic tool to determine the integrity of the spinal cord and the extension response it elicits may seem otherwise useless. In fact, the tendon tap taps into one aspect of a critical building block of mammalian motor control, the stretch reflexes. Stretch reflexes are often thought to quickly resist unexpected changes in muscle length via a very simple circuit in the spinal cord, and this is one circuit that the tendon tap engages. It turns out, however, that stretch reflexes support a myriad of functions and are highly flexible. Under naturalistic conditions, stretch reflexes are shaped by peripheral physiology and engage neural circuits spanning the spinal cord, brainstem and cerebral cortex. In this Primer, we outline what is currently known about stretch reflex function and its underlying mechanisms, with a specific focus on how the cascade of nested responses collectively known as stretch reflexes interact with and build off of one another to support real-world motor behavior.

Voluntary modification of rapid tactile-motor responses during reaching differs from its visuomotor counterpart.

People commonly hold and manipulate a variety of objects in everyday life, and these objects have different physical properties. To successfully control this wide range of objects, people must associate new patterns of tactile stimuli with appropriate motor outputs. We performed a series of experiments investigating the extent to which people can voluntarily modify tactile-motor associations in the context of a rapid tactile-motor response guiding the hand to a moving target (previously described in Pruszynski JA, Johansson RS, Flanagan JR. Curr Biol 26: 788-792, 2016) by using an anti-reach paradigm in which participants were instructed to move their hands in the opposite direction of a target jump. We compared performance to that observed when people make visually guided reaches to a moving target (cf. Day BL, Lyon IN. Exp Brain Res 130: 159-168, 2000; Pisella L, Grea H, Tilikete C, Vighetto A, Desmurget M, Rode G, Boisson D, Rossetti Y. Nat Neurosci 3: 729-736, 2000). When participants had visual feedback, motor responses during the anti-reach task showed early automatic responses toward the moving target before later modification to move in the instructed direction. When the same participants had only tactile feedback, however, they were able to suppress this early phase of the motor response, which occurs <100 ms after the target jump. Our results indicate that while the tactile motor and visual motor systems both support rapid responses that appear similar under some conditions, the circuits underlying responses show sharp distinctions in terms of their malleability.NEW & NOTEWORTHY When people reach toward a visual target that moves suddenly, they automatically correct their reach to follow the object; even when explicitly instructed not to follow a moving visual target, people exhibit an initial incorrect movement before moving in the correct direction. We show that when people use tactile feedback, they do not show an initial incorrect response, even though early muscle activity still occurs.

Effect of Sensory Deprivation on Maximal Force Abilities from Local to Non-local Digits.

The present study investigates the effect of sensory deprivation of the index and middle finger on motor function of all digits during maximal voluntary force production tasks. A total of 27 subjects performed maximal isometric pressing tasks by using different instructed finger combinations. Subjects completed the same tasks in two visits: a control visit when they had normal sensory feedback in all fingers, and an anesthesia visit when digital nerve blocks were performed on their right index and middle fingers. We evaluated three aspects of motor adaptation on both local (anesthetized) and non-local (non-anesthetized) digits during maximal force production: (1) task-relevant and overall force magnitude, (2) force directional application, and (3) digital individuation and force sharing. Our results indicate that selective digital anesthesia resulted in decreased maximal force magnitude, changed direction of force production, and significant changes extended to non-local digits. The motor weakness and inefficiency revealed in the non-local digits implies that sensory information from each digit can be shared across the digits to assist motor execution within the same hand.

Synergic control of action in levodopa-naïve Parkinson's disease patients: I. Multi-finger interaction and coordination.

We explored the origin of the impaired control of action stability in Parkinson's disease (PD) by testing levodopa-naïve PD patients to disambiguate effects of PD from possible effects of long-term exposure to levodopa. Thirteen levodopa-naïve PD patients and 13 controls performed single- and multi-finger force production tasks, including producing a self-paced quick force pulse into a target. A subgroup of patients (n = 10) was re-tested about 1 h after the first dose of levodopa. Compared to controls, PD patients showed lower maximal forces and synergy indices stabilizing total force (reflecting the higher inter-trial variance component affecting total force). In addition, PD patients showed a trend toward shorter anticipatory synergy adjustments (a drop in the synergy index in preparation to a quick action) and larger non-motor equivalent finger force deviations. Lower maximal force, higher unintentional force production (enslaving) and higher inter-trial variance indices occurred in PD patients after one dosage of levodopa. We conclude that impairment in synergies is present in levodopa-naïve patients, mainly in indices reflecting stability (synergy index), but not agility (anticipatory synergy adjustments). A single dose of levodopa, however, did not improve synergy indices, as it did in PD patients on chronic anti-PD medication, suggesting a different mechanism of action. The results suggest that indices of force-stabilizing synergies may be used as an early behavioral sign of PD, although it may not be sensitive to acute drug effects in drug-naïve patients.

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

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

Maintaining arm control during self-triggered and unpredictable unloading perturbations.

We often perform actions where we must break through some resistive force, but want to remain in control during this unpredictable transition; for example, when an object we are pushing on transitions from static to dynamic friction and begins to move. We designed a laboratory task to replicate this situation in which participants actively pushed against a robotic manipulandum until they exceeded an unpredictable threshold, at which point the manipulandum moved freely. Human participants were instructed to either stop the movement of the handle following this unloading perturbation, or to continue pushing. We found that participants were able to modulate their reflexes in response to this unpredictable and self-triggered unloading perturbation according to the instruction they were following, and that this reflex modulation could not be explained by pre-perturbation muscle state. However, in a second task, where participants reactively produced force during the pre-unloading phase in response to the robotic manipulandum to maintain a set hand position, they were unable to modulate their reflexes in the same task-dependent way. This occurred even though the forces they produced were matched to the first task and they had more time to prepare for the unloading event. We suggest this disparity occurs because of different neural circuits involved in posture and movement, meaning that participants in the first task did not require additional time to switch from postural to movement control.

Stability of hand force production. II. Ascending and descending synergies.

We combined the theory of neural control of movement with referent coordinates and the uncontrolled manifold hypothesis to investigate multifinger coordination. We tested hypotheses related to stabilization of performance by covarying control variables, translated into apparent stiffness and referent coordinate, at different levels of an assumed hierarchy of control. Subjects produced an accurate combination of total force and total moment of force with the four fingers under visual feedback on both variables and after feedback was partly or completely removed. The "inverse piano" device was used to estimate control variables. We observed strong synergies in the space of hypothetical control variables that stabilized total force and moment of force, as well as weaker synergies stabilizing individual finger forces; whereas the former were attenuated by alteration of visual feedback, the latter were much less affected. In addition, we investigated the organization of "ascending synergies" stabilizing task-level control variables by covaried adjustments of finger-level control variables. We observed intertrial covariation of individual fingers' referent coordinates that stabilized hand-level referent coordinate, but we observed no such covariation for apparent stiffness. The observations suggest the existence of both descending and ascending synergies in a hierarchical control system. They confirm a trade-off between synergies at different levels of control and corroborate the hypothesis on specialization of different fingers for the control of force and moment. The results provide strong evidence for the importance of central back-coupling loops in ensuring stability of action. NEW & NOTEWORTHY We expand analysis of action in the space of hypothetical control variables to hierarchically organized multieffector systems. We also introduce the novel concept of ascending synergies, which reflect covariation of control variables to individual effectors (fingers) that stabilize task-specific control variables at a hierarchically higher, task-specific level (hand).

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

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

Performance drifts in two-finger cyclical force production tasks performed by one and two actors.

We explored changes in the cyclical two-finger force performance task caused by turning visual feedback off performed either by the index and middle fingers of the dominant hand or by two index fingers of two persons. Based on an earlier study, we expected drifts in finger force amplitude and midpoint without a drift in relative phase. The subjects performed two rhythmical tasks at 1 Hz while paced by an auditory metronome. One of the tasks required cyclical changes in total force magnitude without changes in the sharing of the force between the two fingers. The other task required cyclical changes in the force sharing without changing total force magnitude. Subjects were provided with visual feedback, which showed total force magnitude and force sharing via cursor motion along the vertical and horizontal axes, respectively. Further, visual feedback was turned off, first on the variable that was not required to change and then on both variables. Turning visual feedback off led to a mean force drift toward lower magnitudes while force amplitude increased. There was a consistent drift in the relative phase in the one-hand task with the index finger leading the middle finger. No consistent relative phase drift was seen in the two-person tasks. The shape of the force cycle changed without visual feedback reflected in the lower similarity to a perfect cosine shape and in the higher time spent at lower force magnitudes. The data confirm findings of earlier studies regarding force amplitude and midpoint changes, but falsify predictions of an earlier proposed model with respect to the relative phase changes. We discuss factors that could contribute to the observed relative phase drift in the one-hand tasks including the leader-follower pattern generalized for two-effector tasks performed by one person.

Force illusions and drifts observed during muscle vibration.

We explored predictions of a scheme that views position and force perception as a result of measuring proprioceptive signals within a reference frame set by ongoing efferent process. In particular, this hypothesis predicts force illusions caused by muscle vibration and mediated via changes in both afferent and efferent components of kinesthesia. Healthy subjects performed accurate steady force production tasks by pressing with the four fingers of one hand (the task hand) on individual force sensors with and without visual feedback. At various times during the trials, subjects matched the perceived force using the other hand. High-frequency vibration was applied to one or both of the forearms (over the hand and finger extensors). Without visual feedback, subjects showed a drop in the task hand force, which was significantly smaller under the vibration of that forearm. Force production by the matching hand was consistently higher than that of the task hand. Vibrating one of the forearms affected the matching hand in a manner consistent with the perception of higher magnitude of force produced by the vibrated hand. The findings were consistent between the dominant and nondominant hands. The effects of vibration on both force drift and force mismatching suggest that vibration led to shifts in both signals from proprioceptors and the efferent component of perception, the referent coordinate and/or coactivation command. The observations fit the hypothesis on combined perception of kinematic-kinetic variables with little specificity of different groups of peripheral receptors that all contribute to perception of forces and coordinates. NEW & NOTEWORTHY We show that vibration of hand/finger extensors produces consistent errors in finger force perception. Without visual feedback, finger force drifted to lower values without a drift in the matching force produced by the other hand; hand extensor vibration led to smaller finger force drift. The findings fit the scheme with combined perception of kinematic-kinetic variables and suggest that vibration leads to consistent shifts of the referent coordinate and, possibly, of coactivation command to the effector.

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

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

The recovery response to a novel unannounced laboratory-induced slip: The "first trial effect" in older adults.

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 (≥65year 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.

Unintentional force changes in cyclical tasks performed by an abundant system: Empirical observations and a dynamical model.

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 1Hz, 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.

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

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

Interpersonal synergies: static prehension tasks performed by two actors.

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.

Force-stabilizing synergies in motor tasks involving two actors.

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.

Task-specific stability of multifinger steady-state action.

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.