Reaction of human walking to transient block of vision: analysis in the context of indirect, referent control of motor actions.

Human locomotion may result from monotonic shifts in the referent position, R, of the body in the environment. R is also the spatial threshold at which muscles can be quiescent but are activated depending on the deflection of the current body configuration Q from R. Shifts in R are presumably accomplished with the participation of proprioceptive and visual feedback and responsible for transferring stable body balance (equilibrium) from one place in the environment to another, resulting in rhythmic activity of multiple muscles by a central pattern generator (CPG). We tested predictions of this two-level control scheme. In particular, in response to a transient block of vision during locomotion, the system can temporarily slow shifts in R. As a result, the phase of rhythmical movements of all four limbs will be changed for some time, even though the rhythm and other characteristics of locomotion will be fully restored after perturbation, a phenomenon called long-lasting phase resetting. Another prediction of the control scheme is that the activity of multiple muscles of each leg can be minimized reciprocally at specific phases of the gait cycle both in the presence and absence of vision. Speed of locomotion is related to the rate of shifts in the referent body position in the environment. Results confirmed that human locomotion is likely guided by feedforward shifts in the referent body location, with subsequent changes in the activity of multiple muscles by the CPG. Neural structures responsible for shifts in the referent body configuration causing locomotion are suggested.

Shifts in the eye-centered frame of reference may underlie saccades, visual perception, and eye-hand coordination.

Conventional, computational theories limit the understanding of how action and perception are controlled. In an alternative scheme, the nervous system controls the values of physical and neurophysiological parameters that predetermine the choice of the spatial frames of reference (FRs) for action and perception. For example, all possible eye positions, Q, can be considered as comprising a spatial FR in which extraocular muscles (EOMs) stabilize gaze directions. The origin or referent point of this FR is a specific, threshold eye position, R, at which EOMs can be quiescent but activated depending on the difference between Q and R. Starting before eye motion, shifts in R cause displacement of the FR and resetting of the stable equilibrium position to which the eyes are forced to move. Rather than corollary discharge, the depiction of visual images integrated across the entire retina in the shifted spatial FR is responsible for remapping visual receptive fields and visual constancy. These suggestions are illustrated in computer models of saccades in the referent control framework in humans and monkeys. The existence of three types of visual RF remapping during saccades is suggested. Properly scaled, shifts in the R underlying a saccade are transmitted to motoneurons of arm muscles to guide reach-to-grasp motion in the same, eye-centered FR. Some predictions of the proposed control scheme have been verified and new tests are suggested. The scheme is applicable to several eye-hand coordination deficits including micrography in Parkinson's disease and explains why vision helps deafferented subjects diminish movement deficits.

Eye and head movements and vestibulo-ocular reflex in the context of indirect, referent control of motor actions.

Conventional explanations of the vestibulo-ocular reflex (VOR) and eye and head movements are revisited by considering two alternative frameworks addressing the question of how the brain controls motor actions. Traditionally, biomechanical and/or computational frameworks reflect the views of several prominent scholars of the past, including Helmholtz and von Holst, who assumed that the brain directly specifies the desired motor outcome and uses efference copy to influence perception. However, empirical studies resulting in the theory of referent control of action and perception (an extension of the equilibrium-point hypothesis) revealed that direct specification of motor outcome is inconsistent with nonlinear properties of motoneurons and with the physical principle that the brain can control motor actions only indirectly, by changing or maintaining the values of neurophysiological parameters that influence, but can remain independent of, biomechanical variables. Some parameters are used to shift the origin (referent) points of spatial frames of reference (FRs) or system of coordinates in which motor actions emerge without being predetermined. Parameters are adjusted until the emergent motor actions meet the task demands. Several physiological parameters and spatial FRs have been identified, supporting the notion of indirect, referent control of movements. Instead of integration of velocity-dependent signals, position-dimensional referent signals underlying head motion can likely be transmitted to motoneurons of extraocular muscles. This would produce compensatory eye movement preventing shifts in gaze during head rotation, even after bilateral destruction of the labyrinths. The referent control framework symbolizes a shift in the paradigm for the understanding of VOR and eye and head movement production.

Stability of reaching during standing in stroke.

Reaching from standing requires simultaneous adjustments of focal and postural task elements. We investigated the ability of people with stroke to stabilize the endpoint trajectory while maintaining balance during standing reaches. Nineteen stroke and 11 age-equivalent healthy subjects reached toward a target (n = 30 trials) located beyond arm length from standing. Endpoint and center-of-mass (COM) trajectories were analyzed using the uncontrolled manifold (UCM) approach, with segment angles as elemental variables. A synergy index (SI) represented the normalized difference between segment angle combinations, leading to endpoint or COM trajectory stabilization (VUCM) and lack of stabilization (in an orthogonal space; VORT). A higher SI reflects greater stability. In both groups, the endpoint SI (SIEND) decreased in parallel with endpoint velocity and returned close to baseline at the end of the movement. The range of SIEND was significantly greater in stroke (median: 0.87; QR:0.54) compared with healthy subjects (median: 0.58; QR: 0.33; P = 0.009). In both groups, the lowest SIEND occurred at the endpoint peak velocity, whereas the minimal SIEND of the stroke group (median: 0.51; QR:0.41) was lower than the healthy group (median: 0.25; QR: 0.50; P = 0.033). The COM SI (SICOM) remained stable in both groups (~0.8). The maintenance of a high SICOM despite a large reduction of SIEND in stroke subjects suggests that kinematic redundancy was effectively used to stabilize the COM position, but less so for endpoint position stabilization. Both focal and postural task elements should be considered when analyzing whole body reaching deficits in patients with stroke.NEW & NOTEWORTHY Reaching from standing requires simultaneous adjustments of endpoint and center-of-mass (COM) positions. We used uncontrolled manifold analysis to investigate the impact of stroke on the ability to use kinematic redundancy in this task. Our results showed that COM position was stabilized, whereas endpoint trajectory was more variable in stroke than healthy subjects. Enhancing the capacity to meet multiple task goals may be beneficial for motor recovery after stroke.

Visual deprivation is met with active changes in ground reaction forces to minimize worsening balance and stability during walking.

Previous studies suggest that visual information is essential for balance and stability of locomotion. We investigated whether visual deprivation is met with active reactions tending to minimize worsening balance and stability during walking in humans. We evaluated effects of vision on kinetic characteristics of walking on a treadmill-ground reaction forces (GRFs) and shifts in the center of mass (COM). Young adults (n = 10) walked on a treadmill at a comfortable speed. We measured three orthogonal components of GRFs and COM shifts during no-vision (NV) and full-vision (FV) conditions. We also computed the dynamic balance index (DN)-the perpendicular distance from the projection of center of mass (pCOM) to the inter-foot line (IFL) normalized to half of the foot length. Locally weighted regression smoothing with alpha-adjusted serial T tests was used to compare GRFs and DN between two conditions during the entire stance phase. Results showed significant differences in GRFs between FV and NV conditions in vertical and ML directions. Variability of peak forces of all three components of GRF increased in NV condition. We also observed significant increase in DN for NV condition in eight out of ten subjects. The pCOM was kept within BOS during walking, in both conditions, suggesting that body stability was actively controlled by adjusting three components of GRFs during NV walking to minimize stability loss and preserve balance.

Development of vertical and forward jumping skills in typically developing children in the context of referent control of motor actions.

The empirically based referent control theory of motor actions provides a new framework for understanding locomotor maturation. Mature movement patterns of referent control are characterized by periods of minimization of activity across multiple muscles (global electromyographic [EMG] minima) resulting from transient matching between actual and referent body configurations. We identified whether locomotor maturation in young children was associated with (a) development of referent control and (b) children's frequency of participation in everyday activities evaluated by parents. Kinematics and EMG activity were recorded from typically developing children (n = 15, 3-5 years) and young adults (n = 10, 18-25 years) while walking, vertical or forward jumping. Presence and location of global EMG minima in movement cycles, slopes of ankle vertical/sagittal displacements, and shoulder displacement ratios were evaluated. Children had fewer global EMG minima compared to adults during specific phases of vertical and forward jumps. Ankle displacement profiles for walking and jumping forward were related to each other in adults, whereas those for walking and vertical jumping were related in children. Higher frequency of participation was significantly correlated with more mature jumping patterns in children. A decrease in the number of global EMG minima and changes in ankle movement patterns could be indicators of locomotor immaturity in typically developing children.

Indirect, referent control of motor actions underlies directional tuning of neurons.

Many neurons of the primary motor cortex (M1) are maximally sensitive to "preferred" hand movement directions and generate progressively less activity with movements away from these directions. M1 activity also correlates with other biomechanical variables. These findings are predominantly interpreted in a framework in which the brain preprograms and directly specifies the desired motor outcome. This approach is inconsistent with the empirically derived equilibrium-point hypothesis, in which the brain can control motor actions only indirectly, by changing neurophysiological parameters that may influence, but remain independent of, biomechanical variables. The controversy is resolved on the basis of experimental findings and theoretical analysis of how sensory and central influences are integrated in the presence of the fundamental nonlinearity of neurons: electrical thresholds. In the presence of sensory inputs, electrical thresholds are converted into spatial thresholds that predetermine the position of the body segments at which muscles begin to be activated. Such thresholds may be considered as referent points of respective spatial frames of reference (FRs) in which neurons, including motoneurons, are centrally predetermined to work. By shifting the referent points of respective FRs, the brain elicits intentional actions. Pure involuntary reactions to perturbations are accomplished in motionless FRs. Neurons are primarily sensitive to shifts in referent directions, i.e., shifts in spatial FRs, whereas emergent neural activity may or may not correlate with different biomechanical variables depending on the motor task and external conditions. Indirect, referent control of posture and movement symbolizes a departure from conventional views based on direct preprogramming of the motor outcome.

Referent control of anticipatory grip force during reaching in stroke: an experimental and modeling study.

To evaluate normal and impaired control of anticipatory grip force (GF) modulation, we compared GF production during horizontal arm movements in healthy and post-stroke subjects, and, based on a physiologically feasible dynamic model, determined referent control variables underlying the GF-arm motion coordination in each group. 63% of 13 healthy and 48% of 13 stroke subjects produced low sustained initial force (< 10 N) and increased GF prior to arm movement. Movement-related GF increases were higher during fast compared to self-paced arm extension movements only in the healthy group. Differences in the patterns of anticipatory GF increases before the arm movement onset between groups occurred during fast extension arm movement only. In the stroke group, longer delays between the onset of GF change and elbow motion were related to clinical upper limb deficits. Simulations showed that GFs could emerge from the difference between the actual and the referent hand aperture (Ra) specified by the CNS. Similarly, arm movement could result from changes in the referent elbow position (Re) and could be affected by the co-activation (C) command. A subgroup of stroke subjects, who increased GF before arm movement, could specify different patterns of the referent variables while reproducing the healthy typical pattern of GF-arm coordination. Stroke subjects, who increased GF after arm movement onset, also used different referent strategies than controls. Thus, altered anticipatory GF behavior in stroke subjects may be explained by deficits in referent control.

Vestibular and corticospinal control of human body orientation in the gravitational field.

Body orientation with respect to the direction of gravity changes when we lean forward from upright standing. We tested the hypothesis that during upright standing, the nervous system specifies the referent body orientation that defines spatial thresholds for activation of multiple muscles across the body. To intentionally lean the body forward, the system is postulated to transfer balance and stability to the leaned position by monotonically tilting the referent orientation, thus increasing the activation thresholds of ankle extensors and decreasing their activity. Consequently, the unbalanced gravitational torque would start to lean the body forward. With restretching, ankle extensors would be reactivated and generate increasing electromyographic (EMG) activity until the enhanced gravitational torque would be balanced at a new posture. As predicted, vestibular influences on motoneurons of ankle extensors evaluated by galvanic vestibular stimulation were smaller in the leaned compared with the upright position, despite higher tonic EMG activity. Defacilitation of vestibular influences was also observed during forward leaning when the EMG levels in the upright and leaned position were equalized by compensating the gravitational torque with a load. The vestibular system is involved in the active control of body orientation without directly specifying the motor outcome. Corticospinal influences originating from the primary motor cortex evaluated by transcranial magnetic stimulation remained similar at the two body postures. Thus, in contrast to the vestibular system, the corticospinal system maintains a similar descending facilitation of motoneurons of leg muscles at different body orientations. The study advances the understanding of how body orientation is controlled. NEW & NOTEWORTHY The brain changes the referent body orientation with respect to gravity to lean the body forward. Physiologically, this is achieved by shifts in spatial thresholds for activation of ankle muscles, which involves the vestibular system. Results advance the understanding of how the brain controls body orientation in the gravitational field. The study also extends previous evidence of empirical control of motor function, i.e., without the reliance on model-based computations and direct specification of motor outcome.

Spasticity may obscure motor learning ability after stroke.

Previous motor learning studies based on adapting movements of the hemiparetic arm in stroke subjects have not accounted for spasticity occurring in specific joint ranges (spasticity zones), resulting in equivocal conclusions about learning capacity. We compared the ability of participants with stroke to rapidly adapt elbow extension movements to changing external load conditions outside and inside spasticity zones. Participants with stroke ( n = 12, aged 57.8 ± 9.6 yr) and healthy age-matched controls ( n = 8, 63.5 ± 9.1 yr) made rapid 40°-50° horizontal elbow extension movements from an initial (3°) to a final (6°) target. Sixteen blocks (6-10 trials/block) consisting of alternating loaded (30% maximal voluntary contraction) and nonloaded trials were made in one (controls) or two sessions (stroke; 1 wk apart). For the stroke group, the tonic stretch reflex threshold angle at which elbow flexors began to be activated during passive elbow extension was used to identify the beginning of the spasticity zone. The task was repeated in joint ranges that did or did not include the spasticity zone. Error correction strategies were identified by the angular positions before correction and compared between groups and sessions. Changes in load condition from no load to load and vice versa resulted in undershoot and overshoot errors, respectively. Stroke subjects corrected errors in 1-4 trials compared with 1-2 trials in controls. When movements did not include the spasticity zone, there was an immediate decrease in the number of trials needed to restore accuracy, suggesting that the capacity to learn may be preserved after stroke but masked by the presence of spasticity. NEW & NOTEWORTHY When arm movements were made outside, instead of inside, the range affected by spasticity, there was an immediate decrease in the number of trials needed to restore accuracy in response to a change in the external load. This suggests that motor learning processes may be preserved in patients with stroke but masked by the presence of spasticity in specific joint ranges. This has important implications for designing rehabilitation interventions predicated on motor learning principles.

Referent control of the orientation of posture and movement in the gravitational field.

This study addresses the question of how posture and movement are oriented with respect to the direction of gravity. It is suggested that neural control levels coordinate spatial thresholds at which multiple muscles begin to be activated to specify a referent body orientation (RO) at which muscle activity is minimized. Under the influence of gravity, the body is deflected from the RO to an actual orientation (AO) until the emerging muscle activity and forces begin to balance gravitational forces and maintain body stability. We assumed that (1) during quiet standing on differently tilted surfaces, the same RO and thus AO can be maintained by adjusting activation thresholds of ankle muscles according to the surface tilt angle; (2) intentional forward body leaning results from monotonic ramp-and-hold shifts in the RO; (3) rhythmic oscillation of the RO about the ankle joints during standing results in body swaying. At certain sway phases, the AO and RO may transiently overlap, resulting in minima in the activity of multiple muscles across the body. EMG kinematic patterns of the 3 tasks were recorded and explained based on the RO concept that implies that these patterns emerge due to referent control without being pre-programmed. We also confirmed the predicted occurrence of minima in the activity of multiple muscles at specific body configurations during swaying. Results re-affirm previous rejections of model-based computational theories of motor control. The role of different descending systems in the referent control of posture and movement in the gravitational field is considered.

Threshold position control of anticipation in humans: a possible role of corticospinal influences.

KEY POINTS: Sudden unloading of preloaded wrist muscles elicits motion to a new wrist position. Such motion is prevented if subjects unload muscles using the contralateral arm (self-unloading). Corticospinal influences originated from the primary motor cortex maintain tonic influences on motoneurons of wrist muscles before sudden unloading but modify these influences prior to the onset and until the end of self-unloading. Results are interpreted based on the previous finding that intentional actions are caused by central, particularly corticospinal, shifts in the spatial thresholds at which wrist motoneurons are activated, thus predetermining the attractor point at which the neuromuscular periphery achieves mechanical balance with environment forces. By maintaining or shifting the thresholds, descending systems let body segments go to the equilibrium position in the respective unloading tasks without the pre-programming of kinematics or muscle activation patterns. The study advances the understanding of how motor actions in general, and anticipation in particular, are controlled. ABSTRACT: The role of corticospinal (CS) pathways in anticipatory motor actions was evaluated using transcranial magnetic stimulation (TMS) of the primary motor cortex projecting to motoneurons (MNs) of wrist muscles. Preloaded wrist flexors were suddenly unloaded by the experimenter or by the subject using the other hand (self-unloading). After sudden unloading, the wrist joint involuntarily flexed to a new position. In contrast, during self-unloading the wrist remained almost motionless, implying that an anticipatory postural adjustment occurred. In the self-unloading task, anticipation was manifested by a decrease in descending facilitation of pre-activated flexor MNs starting ∼72 ms before changes in the background EMG activity. Descending facilitation of extensor MNs began to increase ∼61 ms later. Conversely, these influences remained unchanged before sudden unloading, implying the absence of anticipation. We also tested TMS responses during EMG silent periods produced by brief muscle shortening, transiently resulting in similar EMG levels before the onset and after the end of self-unloading. We found reduced descending facilitation of flexor MNs after self-unloading. To explain why the wrist excursion was minimized in self-unloading due to these changes in descending influences, we relied on previous demonstrations that descending systems pre-set the threshold positions of body segments at which muscles begin to be activated, thus predetermining the equilibrium point to which the system is attracted. Based on this notion, a more consistent explanation of the kinematic, EMG and descending patterns in the two types of unloading is proposed compared to the alternative notion of direct pre-programming of kinematic and/or EMG patterns.

Referent control and motor equivalence of reaching from standing.

Motor actions may result from central changes in the referent body configuration, defined as the body posture at which muscles begin to be activated or deactivated. The actual body configuration deviates from the referent configuration, particularly because of body inertia and environmental forces. Within these constraints, the system tends to minimize the difference between these configurations. For pointing movement, this strategy can be expressed as the tendency to minimize the difference between the referent trajectory (RT) and actual trajectory (QT) of the effector (hand). This process may underlie motor equivalent behavior that maintains the pointing trajectory regardless of the number of body segments involved. We tested the hypothesis that the minimization process is used to produce pointing in standing subjects. With eyes closed, 10 subjects reached from a standing position to a remembered target located beyond arm length. In randomly chosen trials, hip flexion was unexpectedly prevented, forcing subjects to take a step during pointing to prevent falling. The task was repeated when subjects were instructed to intentionally take a step during pointing. In most cases, reaching accuracy and trajectory curvature were preserved due to adaptive condition-specific changes in interjoint coordination. Results suggest that referent control and the minimization process associated with it may underlie motor equivalence in pointing. NEW & NOTEWORTHY: Motor actions may result from minimization of the deflection of the actual body configuration from the centrally specified referent body configuration, in the limits of neuromuscular and environmental constraints. The minimization process may maintain reaching trajectory and accuracy regardless of the number of body segments involved (motor equivalence), as confirmed in this study of reaching from standing in young healthy individuals. Results suggest that the referent control process may underlie motor equivalence in reaching.

Active sensing without efference copy: referent control of perception.

Although action and perception are different behaviors, they are likely to be interrelated, as implied by the notions of perception-action coupling and active sensing. Traditionally, it has been assumed that the nervous system directly preprograms motor commands required for actions and uses a copy of them called efference copy (EC) to also influence our senses. This review offers a critical analysis of the EC concept by identifying its limitations. An alternative to the EC concept is based on the experimentally confirmed notion that sensory signals from receptors are perceived relative to referent signals specified by the brain. These referents also underlie the control of motor actions by predetermining where, in the spatial domain, muscles can work without preprogramming how they should work in terms of motor commands or EC. This approach helps solve several problems of action and explain several sensory experiences, including position sense and the sense that the world remains stationary despite changes in its retinal image during eye or body motion (visual space constancy). The phantom limb phenomenon and other kinesthetic illusions are also explained within this framework.

Implicit learning and generalization of stretch response modulation in humans.

Adaptation of neural responses to repeated muscle stretching likely represents implicit learning to minimize muscle resistance to perturbations. To test this hypothesis, the forearm was placed on a horizontal manipulandum. Elbow flexors or extensors compensated an external load and were stretched by 20° or 70° rotations. Participants were instructed not to intervene by intentionally modifying the muscle resistance elicited by stretching. In addition to phasic stretch reflexes (SRs), muscle stretching was associated with inhibitory periods (IPs) in the ongoing muscle activity starting at minimal latencies of ∼35 ms. The SR amplitude decreased dramatically across 5-12 trials and was not restored after a resting period of 3-5 min, despite the increase in stretch amplitude from 20° to 70°, but IPs remained present. When SRs were suppressed, stretching of originally nonstretched, antagonist muscles initiated after the rest period showed immediate SR suppression while IPs remained present in the first and subsequent trials. Adaptation to muscle stretching thus includes features characteristic of implicit learning such as memory consolidation and generalization. Adaptation may be achieved by central shifts in the threshold positions at which muscles begin to be activated. Shifts are thought to be prepared in advance and triggered with stretch onset. Threshold position resetting provides a comprehensive explanation of the results in the broader context of the control of posture, movement, and motor learning in the healthy and damaged nervous system.

The Relationship Between Postural and Movement Stability.

Postural stabilization is provided by stretch reflexes, intermuscular reflexes, and intrinsic muscle properties. Taken together, these posture-stabilizing mechanisms resist deflections from the posture at which balance of muscle and external forces is maintained. Empirical findings suggest that for each muscle, these mechanisms become functional at a specific, spatial threshold-the muscle length or respective joint angle at which motor units begin to be recruited. Empirical data suggest that spinal and supraspinal centers can shift the spatial thresholds for a group of muscles that stabilized the initial posture. As a consequence, the same stabilizing mechanisms, instead of resisting motion from the initial posture, drive the body to another stable posture. In other words by shifting spatial thresholds, the nervous system converts movement resisting to movement-producing mechanisms. It is illustrated that, contrary to conventional view, this control strategy allows the system to transfer body balance to produce locomotion and other actions without loosing stability at any point of them. It also helps orient posture and movement with the direction of gravity. It is concluded that postural and movement stability is provided by a common mechanism.

Referent Control of Side-to-Side Body-Weight Transfer During Standing and Stepping in Adults.

Research suggests that locomotion may be primarily caused by shifting stable body balance from one location in the environment to another with subsequent rhythmical muscle activation by the central pattern generator (CPG), constituting a multi-level control system. All levels interact with environmental forces affected by proprioceptive and vestibular reflexes as well as vision. A similar multi-level control schema is likely used to shift body balance laterally when the body weight is rhythmically transferred from side-to-side. In order to do so, the system shifts a specific body posture in space. This body posture is referred to as the threshold or referent body posture, R, at which all muscles involved can be at rest but are activated depending on the deflection of the actual body posture, Q, from R. This concept has previously been investigated for forward and backward locomotion. The purpose of the present study was to verify if it was also applicable to locomotor tasks in other directions such as sidestepping. We predicted that during sidestepping, the actual and referent posture can transiently match each other bringing the activity of multiple muscles to a minimum. The existence of such minima was demonstrated in healthy adults performing three locomotor tasks involving shifts of the body weight from side-to-side thus further supporting the validity of the multi-level control scheme of locomotion.

Altered Anticipatory Postural Adjustments During Whole-Body Reaching in Subjects With Stroke.

BACKGROUND: Coordination between arm movements and postural adjustments is crucial for reaching-while-stepping tasks involving both anticipatory postural adjustments (APAs) and compensatory movements to effectively propel the whole-body forward so that the hand can reach the target. Stroke impairs the ability to coordinate the action of multiple body segments but the underlying mechanisms are unclear. Objective. To determine the effects of stroke on reaching performance and APAs during whole-body reaching. METHODS: We tested arm reaching in standing (stand-reach) and reaching-while-stepping (step-reach; 15 trials/condition) in individuals with chronic stroke (n = 18) and age-matched healthy subjects (n = 13). Whole-body kinematics and kinetic data were collected during the tasks. The primary outcome measure for step-reach was "gain" (g), defined as the extent to which the hip displacement contributing to hand motion was neutralized by appropriate changes in upper limb movements (g = 1 indicates complete compensation) and APAs measured as spatio-temporal profiles of the center-of-pressure shifts preceding stepping. RESULTS: Individuals with stroke had lower gains and altered APAs compared to healthy controls. In addition, step onset was delayed, and the timing of endpoint, trunk, and foot movement offset was prolonged during step-reach compared to healthy controls. Those with milder sensorimotor impairment and better balance function had higher gains. Altered APAs were also related to reduced balance function. CONCLUSIONS: Altered APAs and prolonged movement offset in stroke may lead to a greater reliance on compensatory arm movements. Altered APAs in individuals with stroke may be associated with a reduced shift of referent body configuration during the movement.

Reduced gait stability in high-functioning poststroke individuals.

Falls during walking are a major cause of poststroke injury, and walking faster may decrease the ability to recover following a gait perturbation. We compared gait stability between high-functioning poststroke individuals and controls and evaluated the effect of gait speed on gait stability. Ten stroke subjects and ten age-matched controls walked on a self-paced treadmill at two speeds (matched/faster). Movement of the nonparetic/dominant leg was arrested unexpectedly at early swing. Poststroke individuals lowered the perturbed leg following perturbation (58% of cases) while controls maintained the leg elevated (49% of cases; P < 0.01). In poststroke individuals, double-support duration was restored later than in controls (4.6 ± 0.8 vs. 3.2 ± 0.3 strides; P < 0.007), and long-term phase shifts of arm and leg movements were larger and less coordinated on the paretic side. A moderate speed increase (~20%) enhanced the incidence of leg lowering in controls but not in stroke subjects. Faster walkers in both groups had a more coordinated response, limited to the nonparetic side in the stroke group. However, faster walkers were not more stable following perturbation. Our results suggest that gait perturbations can target basic control processes and identify neurological locomotor deficits in individuals with fall risk. Central regulation of body translation in space is involved in recovery of steady-state walking. Impaired descending control (stroke) decreases the ability of the motor system to recover from perturbations and regulate interlimb phase relationships, especially when changing gait speed. However, interlimb coordination may not be a major factor in the recovery of gait stability.