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Background Motor performance is a complex process controlled in task-specific spatial frames of reference (FRs). Movements can be made within the framework of the body (egocentric FR) or external space (exocentric FR). People with stroke have impaired reaching, which may be related to deficits in movement production in different FRs. Objective To characterize rapid motor responses to changes in the number of degrees of freedom for movements made in different FRs and their relationship with sensorimotor and cognitive impairment in individuals with mild chronic stroke. Methods Healthy and poststroke individuals moved their hand along the contralateral forearm (egocentric task) and between targets in the peripersonal space (exocentric task) without vision while flexing the trunk. Trunk movement was blocked in randomized trials. Results For the egocentric task, controls produced the same endpoint trajectories in both conditions (free- and blocked-trunk) by preserving similar shoulder-elbow interjoint coordination (IJC). However, endpoint trajectories were dissimilar because of altered IJC in stroke. For the exocentric task, controls produced the same endpoint trajectories when the trunk was free or blocked by rapidly changing the IJC, whereas this was not the case in stroke. Deficits in exocentric movement after stroke were related to cognitive but not sensorimotor impairment. Conclusions Individuals with mild stroke have deficits rapidly responding to changing conditions for complex reaching tasks. This may be related to cognitive deficits and limitations in the regulation of tonic stretch reflex thresholds. Such deficits should be considered in rehabilitation programs encouraging the reintegration of the affected arm into activities of daily living.

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.

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.

There are contralateral and less studied ipsilateral (i), indirect cortical descending projections to motoneurons (MNs). We compared ipsilateral cortical descending influences on MNs of wrist flexors by applying transcranial magnetic stimulation (TMS) over the right primary motor cortex at actively maintained flexion and extension wrist positions in uni- and bimanual tasks in right-handed participants (n = 23). The iTMS response includes a short latency (~ 25 ms) motor evoked potential (iMEP), a silent period (iSP) and a long latency (~ 60 ms) facilitation called rebound (iRB). We also investigated whether the interaction between the two hands while holding an object in a bimanual task involves ipsilateral cortical descending influences. In the unimanual task, iTMS responses in the right wrist flexors were unaffected by changes in wrist position. In the bimanual task with an object, iMEPs in the right wrist flexors were larger when the ipsilateral wrist was in flexion compared to extension. Without the object, only iRB were larger when the ipsilateral wrist was extended. Thus, ipsilateral cortical descending influences on MNs were modulated only in bimanual tasks and depended on how the two hands interacted. It is concluded that the left and right cortices cooperate in bimanual tasks involving holding an object with both hands, with possible involvement of oligo- and poly-synaptic, as well as transcallosal projections to MNs. The possible involvement of spinal and transcortical stretch and cutaneous reflexes in bimanual tasks when holding an object is discussed in the context of the well-established notion that indirect, referent control underlies motor actions.

Objectives The corticospinal system (CS) regulates muscle activation through shifts in muscle-level tonic stretch-reflex thresholds (TSRT). This ability is impaired in stroke and contributes to sensorimotor impairments such as spasticity. We determined the role of CS in elbow flexor activity regulation in healthy and post-stroke subjects. We also determined whether CS modulation deficits were related to sensorimotor impairment intensity in post-stroke individuals. Methods Seventeen healthy (59.8 ± 12.2 yr) and 27 stroke subjects (58.7 ± 10.1 yr) had transcranial magnetic stimulation (TMS) applied over the primary motor cortex (M1) flexor representation to elicit motor-evoked potentials (MEPs) in elbow flexors in different angular positions. In a subset of post-stroke subjects (n = 12), flexor TSRTs were measured in passive and active conditions, and TSRT modulation was determined. Results Position-related MEP amplitude modulation was similar in healthy and mild stroke subjects, while subjects with more severe stroke exhibited less consistent modulation. MEP modulation in stroke was related to clinical upper limb motor impairment, spasticity, and the ability to modulate TSRTs between passive and active elbow movements. Conclusions CS output was closely related to TSRT modulation. Impairments in TSRT regulation may underlie motor deficits in moderate-to-severe post-stroke individuals. Significance Translation of these neurophysiological findings to clinical applications may enhance post-stroke motor recovery.

Objectives Deficits in regulation of tonic stretch reflex thresholds (TSRTs) after stroke occur in elbow flexors and extensors leading to spasticity in specific joint ranges. Threshold deregulation may also be responsible for other deficits such as abnormal activation of passively shortening muscles. Goals were to characterize activation of shortening elbow extensors during passive elbow flexor stretch in individuals with stroke, and identify its relationship to upper-limb motor impairment. Methods Thirty-three participants with unilateral stroke participated. TSRTs in elbow flexors were measured by stretching passive elbow flexors at different velocities. EMG responses were recorded from stretched agonist (biceps) and shortened antagonist (triceps) muscles. Results Triceps activation during passive biceps stretch occurred in all but 4 participants simultaneously with, before or after biceps activation onset. Biceps and triceps activation onsets and durations decreased with stretch velocity. Biceps TSRT and triceps activation magnitude did not correlate with sensorimotor impairment but greater stroke chronicity tended to be related to higher biceps TSRTs (r = 0.406, p = 0.041). Conclusions Stroke may result in both limitations in reciprocal inhibition and excessive agonist-antagonist co-activation, likely from deficits in TSRT modulation in both muscle groups. Significance Since both reciprocal inhibition and co-activation are fundamental to normal motor control, their cooperative action should be considered in designing interventions to increase the ranges of regulation of TSRTs in flexors and extensors to enhance upper limb functional recovery.

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.

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.

International audience; 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.

Objectives Voluntary movements post-stroke are affected by abnormal muscle activation due to exaggerated stretch reflexes (SRs). We examined the ability of post-stroke subjects to regulate SRs in spastic muscles. Methods Elbow flexor and extensor EMGs and joint angle were recorded in 13 subjects with chronic post-stroke spasticity. Muscles were either stretched passively (relaxed arm) or actively (antagonist contraction) at different velocities. Velocity-dependent SR thresholds were defined as angles where stretched muscle EMG exceeded 3SDs of baseline. Sensitivity of SRs to stretch velocity was defined as µ. The regression through thresholds was interpolated to zero velocity to obtain the tonic SR threshold (TSRT) angle. Results Compared to passive stretches, TSRTs during active motion occurred at longer muscle lengths (i.e., increased in flexors and decreased in extensors by 10–40°). Values of μ increased by 1.5–4.0. Changes in flexor TSRTs during active compared to passive stretches were correlated with clinical spasticity (r = −0.68) and arm motor impairment (r = 0.81). Conclusions Spasticity thresholds measured at rest were modulated during active movement. Arm motor impairments were related to the ability to modulate SR thresholds between the two states rather than to passive-state values. Significance Relationship between spasticity and movement disorders may be explained by deficits in SR threshold range of regulation and modifiability, representing a measure of stroke-related sensorimotor deficits.

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 pre-determining 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. This article is protected by copyright. All rights reserved

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.

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.

Control of reflexes is usually associated with central modulation of their sensitivity (gain) or phase-dependent inhibition and facilitation of their influences on motoneurons (reflex gating). Accumulated empirical findings show that the gain modulation and reflex gating are secondary, emergent properties of central control of spatial thresholds at which reflexes become functional. In this way, the system pre-determines, in a feedforward and task-specific way, where, in a spatial domain or a frame of reference, muscles are allowed to work without directly prescribing EMG activity and forces. This control strategy is illustrated by considering reflex adaptation to repeated muscle stretches in healthy subjects, a process associated with implicit learning and generalization. It has also been shown that spasticity, rigidity, weakness and other neurological motor deficits may have a common source – limitations in the range of spatial threshold control elicited by neural lesions.

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 (

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.

Many falls in older adults occur during walking following trips. Following a trip, older adults take longer than younger adults to recover steady-state walking. Although faster gait speed may improve interlimb coordination, it may also increase fall risk in older adults. We hypothesized that older adults would take longer than younger adults to recover from an unexpected perturbation during gait especially when walking faster. Twelve younger (26.3 ± 4.4 years) and 12 older adults (68.5 ± 3.4 years) walked at comfortable, faster and slower speeds when movement of the dominant leg was unexpectedly arrested for 250 ms at 20% swing length. Gait stability was evaluated using the short- and longer-term response to perturbation. In both groups, walking faster diminished the occurrence of elevation and increased that of leg lowering. Older adults took longer than younger adults to recover steady-state walking at all speeds (3.36 ± 0.11 vs. 2.89 ± 0.08 strides) but longer-term recovery of gait stability was not related to gait speed. Arm-leg and inter-arm coordination improved with increasing gait speed in both groups, but older adults had weaker inter-leg coupling following perturbation at all speeds. Although both younger and older adults used speed appropriate responses immediately following perturbation, longer duration of recovery of steady-state walking in older adults may increase fall risk in uncontrolled situations, regardless of gait speed. Recovery from perturbation when walking faster was associated with better interlimb coordination, but not with better gait stability. This indicates that interlimb coordination and gait stability may be distinct features of locomotion.

Background. By involving additional degrees of freedom, the nervous system may preserve hand trajectories when making pointing movements with or without trunk displacement. Previous studies indicate that the potential contribution of trunk movement to hand displacement for movements made within arm reach is neutralized by appropriate compensatory shoulder and elbow rotations. For beyond-the-reach movements, compensatory coordination is attenuated after the hand peak velocity, allowing trunk movement to contribute to hand displacement. Objective. To investigate if the timing and spatial coordination of arm and trunk movements during beyond-the-reach movements is preserved in stroke. Methods. Eleven healthy control subjects and 11 individuals with mild-to-moderate chronic unilateral hemiparesis participated. Arm and trunk kinematics during 60 target reaches to an ipsilaterally placed target were recorded. In 30% of randomly chosen trials, trunk movement was unexpectedly prevented (blocked-trunk trials) by an electromagnetic device, resulting in divergence of the hand trajectory from that in free-trunk trials. Hand trajectories and elbow–shoulder interjoint coordination were compared between trials. Results. In stroke participants, hand trajectory divergence occurred at a shorter movement extent and interjoint coordination patterns diverged at a relatively greater distance compared to controls. Thus, arm movements in stroke participants only partially compensated trunk displacement resulting in the trunk movement contributing to arm movement earlier and to a larger extent during reaching. Conclusion. Individuals with mild-to-moderate stroke have deficits in timing and spatial coordination of arm and trunk movements during different parts of a reaching movement. This deficit may be targeted in therapy to improve upper limb function.

Objective Muscle spasticity following stroke has been shown to result from limitations in the range of regulation of the tonic reflex spatial threshold (ST), i.e., the joint angle at which the stretch reflex begins to act due to descending and segmental influences on motoneurons. The purpose of this study was to determine whether spasticity due to stroke and rigidity due to parkinsonism can be discriminated based on the ST measure. Methods Elbow muscles were stretched at different velocities in healthy, stroke (spasticity) and parkinsonism (rigidity) subjects. The elbow angle at which muscle activation began for each stretch velocity (dynamic ST) and the velocity sensitivity of the ST were measured. Dynamic ST values extrapolated to zero velocity defined the tonic ST. Results Compared to healthy subjects, spasticity and rigidity were associated with a decrease in the range of central regulation of tonic STs. STs were hypersensitive in spastic muscles and either hypo- or inversely sensitive to stretch velocity in rigid muscles. Conclusions ST characteristics discriminate between neurological deficits of muscle tone. Significance Results suggest that spasticity and rigidity result from deficits in descending facilitatory control combined with deficits in dynamic fusimotor or/and presynaptic control of Ia inputs to motoneurons.

The difference between voluntary and involuntary motor actions has been recognized since ancient times, but the nature of this difference remains unclear. We compared corticospinal influences at wrist positions established before and after voluntary motion with those established before and after involuntary motion elicited by sudden removal of a load (the unloading reflex). To minimize the effect of motoneuronal excitability on the evaluation of corticospinal influences, motor potentials from transcranial magnetic stimulation of the wrist motor cortex area were evoked during an EMG silent period produced by brief muscle shortening. The motoneuronal excitability was thus equalized at different wrist positions. Results showed that the unloading reflex was generated in the presence of a corticospinal drive, rather than autonomously by the spinal cord. Although the tonic EMG levels were substantially different, the corticospinal influences remained the same at the pre- and post-unloading wrist positions. These influences however changed when subjects voluntarily moved the wrist to another position. Previous studies showed that the corticospinal system sets the referent position (R) at which neuromuscular posture-stabilizing mechanisms begin to act. In self-initiated actions, the corticospinal system shifts the R to relay these mechanisms to a new posture, thus converting them from mechanisms resisting to those assisting motion. This solves the classical posture-movement problem. In contrast, by maintaining the R value constant, the corticospinal system relies on these posture-stabilizing mechanisms to allow involuntary responses to occur after unloading. Thus, central control strategies underlying the two types of motor actions are fundamentally different.

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.

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.

Most falls in older adults occur when walking, specifically following a trip. This study investigated the short- and longer term responses of young ( n = 24, 27.6 ± 4.5 yr) and older adults ( n = 18, 69.1 ± 4.2 yr) to a trip during gait at comfortable speed and the role of interlimb coordination in recovery from tripping. Subjects walked on a self-paced treadmill when forward movement of their dominant leg was unexpectedly arrested for 250 ms. Recovery of center of mass (COM) movements and of double-support duration following perturbation was determined. In addition, the disruption and recovery of interlimb coordination of the arms and legs was evaluated. Although young and older subjects used similar lower limb strategies in response to the trip, older adults had less stable COM movement patterns before perturbation, had longer transient destabilization (>25%) after perturbation, required more gait cycles to recover double-support duration (older, 3.48 ± 0.7 cycles; young, 2.88 ± 0.4 cycles), and had larger phase shifts that persisted after perturbation (older, −83° to −90°; young, −39° to −42°). Older adults also had larger disruptions to interlimb coordination of the arms and legs. The timing of the initial disruption in coordination was correlated with the disturbance in gait stability only in young adults. In older adults, greater initial COM instability was related to greater longer term arm incoordination. These results suggest a relationship between interlimb coordination and gait stability, which may be associated with fall risk in older adults. Reduced coordination and gait stability suggest a need for stability-related functional training even in high-functioning older adults.

It has previously been established that muscles become active in response to deviations from a threshold (referent) position of the body or its segments, and that intentional motor actions result from central shifts in the referent position. We tested the hypothesis that corticospinal pathways are involved in threshold position control during intentional changes in the wrist position in humans. Subjects moved the wrist from an initial extended to a final flexed position (and vice versa). Passive wrist muscle forces were compensated with a torque motor such that wrist muscle activity was equalized at the two positions. It appeared that motoneuronal excitability tested by brief muscle stretches was also similar at these positions. Responses to mechanical perturbations before and after movement showed that the wrist threshold position was reset when voluntary changes in the joint angle were made. Although the excitability of motoneurons was similar at the two positions, the same transcranial magnetic stimulus (TMS) elicited a wrist extensor jerk in the extension position and a flexor jerk in the flexion position. Extensor motor-evoked potentials (MEPs) elicited by TMS at the wrist extension position were substantially bigger compared to those at the flexion position and vice versa for flexor MEPs. MEPs were substantially reduced when subjects fully relaxed wrist muscles and the wrist was held passively in each position. Results suggest that the corticospinal pathway, possibly with other descending pathways, participates in threshold position control, a process that pre-determines the spatial frame of reference in which the neuromuscular periphery is constrained to work. This control strategy would underlie not only intentional changes in the joint position, but also muscle relaxation. The notion that the motor cortex may control motor actions by shifting spatial frames of reference opens a new avenue in the analysis and understanding of brain function.

Fatigue affects the capacity of muscles to generate forces and is associated with characteristic changes in EMG signals. It may also influence interjoint and intermuscular coordination. To understand better the global effects of fatigue on multijoint movement, we studied movement kinematics and EMG changes in healthy volunteers asked to hammer repetitively. Movement kinematics and the activity of 20 muscles of the arm, trunk, and leg were recorded before and after subjects became fatigued (as measured using a Borg scale). When fatigue was reached, maximal grip strength and elbow range of motion decreased while the EMG amplitude of the contralateral external oblique muscle was increased. Fatigue did not affect shoulder and wrist kinematics or movement frequency. Results suggest that fatigue influences motion at both local and global levels. Specifically, interjoint and intermuscular coordination adapt to compensate for local effects of fatigue and to maintain key movement characteristics, such as the trajectory of the end effector and the movement frequency. Nonlocal compensations may be a focus of future studies of how fatigue affects complex movements such as those typically performed in the workplace.

The grip force holding an object between fingers usually increases before or simultaneously with arm movement thus preventing the object from sliding. We experimentally analyzed and simulated this anticipatory behavior based on the following notions. (1) To move the arm to a new position, the nervous system shifts the threshold position at which arm muscles begin to be recruited. Deviated from their activation thresholds, arm muscles generate activity and forces that tend to minimize this deviation by bringing the arm to a new position. (2) To produce a grip force, with or without arm motion, the nervous system changes the threshold configuration of the hand. This process defines a threshold (referent) aperture (R(a)) of appropriate fingers. The actual aperture (Q(a)) is constrained by the size of the object held between the fingers whereas, in referent position R(a), the fingers virtually penetrate the object. Deviated by the object from their thresholds of activation, hand muscles generate activity and grip forces in proportion to the gap between the Q(a) and R(a). Thus, grip force emerges since the object prevents the fingers from reaching the referent position. (3) From previous experiences, the system knows that objects tend to slide off the fingers when arm movements are made and, to prevent sliding, it starts narrowing the referent aperture simultaneously with or somewhat before the onset of changes in the referent arm position. (4) The interaction between the fingers and the object is accomplished via the elastic pads on the tips of fingers. The pads are compressed not only due to the grip force but also due to the tangential inertial force ("load") acting from the object on the pads along the arm trajectory. Compressed by the load force, the pads move back and forth in the gap between the finger bones and object, thus inevitably changing the normal component of the grip force, in synchrony with and in proportion to the load force. Based on these notions, we simulated experimental elbow movements and grip forces when subjects rapidly changed the elbow angle while holding an object between the index finger and the thumb. It is concluded that the anticipatory increase in the grip force with or without correlation with the tangential load during arm motion can be explained in neurophysiological and biomechanical terms without relying on programming of grip force based on an internal model.

In this chapter, I will describe experiments in humans and animals supporting the notion that the nervous system exercises parametric control of motor actions (see Chap. 2). It will be demonstrated that by changing task-specific parameters, the system allows physical laws to take their course to generate motor outcome without pre-programming of EMG patterns, muscle forces and kinematics. Data will be shown indicating that the nervous system exercises a specific form of parametric control by stipulating spatial thresholds for motoneuronal recruitment, i.e. the threshold muscle lengths or respective joint angles at which α-MNs begin to be activated. In this way, neural control levels pre-determine where, in the spatial domain, neuromuscular elements can work without prescribing how they should work. MNs are recruited or de-recruited depending on the difference between the actual and the threshold muscle lengths, the latter also being dependent on the rate of change in the muscle length.

We addressed the fundamental questions of which variables underlie the control of arm movement and how they are stored in motor memory, reproduced and modified in the process of adaptation to changing load conditions. Such variables are defined differently in two major theories of motor control (internal models and threshold control). To resolve the controversy, these theories were tested (experiment 1) based on their ability to explain why active movement away from a stable posture is not opposed by stabilizing mechanisms (the posture-movement problem). The internal model theory suggests that the system counteracts the opposing forces by increasing the muscle activity in proportion to the distance from the initial posture (position-dependent EMG control). In contrast, threshold control fully excludes these opposing forces by shifting muscle activation thresholds and thus resetting the stabilizing mechanisms to a new posture. Subjects were sitting, holding the vertical handle of a double-joint manipulandum with their right hand and were facing a computer screen on which the handle and target to be reached were displayed. In response to an auditory signal, subjects quickly moved the handle from an initial position to one of two (frontal and sagittal) targets. No load was applied during the movement but in separate trials, a brief perturbation was applied to the handle by torque motors controlling the manipulandum. Perturbations were applied prior to or 3 s after movement offset, in the latter case in one of eight directions. The EMG activity of the majority of the seven recorded muscles was at zero level before movement onset and returned to zero level after movement offset. Those muscles that remained active before or after the movement could be made silent whereas previously silent muscles could be activated after a small passive displacement (several millimeters) elicited by perturbations in appropriate directions. Results showed that the activation thresholds of motoneurons of arm muscles were reset from the initial to a final position and that EMG activity was not position-dependent. These results were inconsistent with the internal model theory but confirmed the threshold control theory. Then the ability of threshold control theory to explain rapid movement adaptation to a position-dependent load was investigated (experiment 2 and 3). Subjects produced fast movement to the frontal target with and without a position-dependent load applied to the handle. Trials were organized in blocks alternating between the load and no-load condition (20 blocks in total, with randomly chosen number of five to ten trials in each). Subjects were instructed "do not correct" in experiment 2 and "correct" movement errors during the trial in experiment 3. Five threshold arm configurations underlying the movement production and adaptation were identified. When instructed "do not correct", movement precision was fully restored on average after two trials. No significant improvement was observed as the experiment progressed despite the fact that the same load condition was repeated after one block of trials. Thus, in each block, the adaptation was made anew, implying that subjects relied on short-term memory and could not recall the threshold arm configurations they specified to accurately reach the same target in the same load condition in previous blocks. When instructed to "correct" within each trial, precision was restored faster, on average after one trial. Major aspects of the production and adaptation of arm movement (including the kinematics, movement errors, instruction-dependent behavior, and absence of position-related EMG activity) are explained in terms of threshold control.

It is usually assumed that proprioceptive feedback comes to motoneurons too late to contribute to the initial activity of agonist muscles during fast arm movements, leading to the suggestion that this feedback is only efficient in slow movements and postural control. The argument does not take into account that the changes in the motoneuronal membrane potentials and the associated changes in the state of spinal neurons preceding the initial activity of muscles deeply affect, in a forward way, the state of reflex systems by shifting their thresholds, as suggested in the lambda model for motor control. As a result, the initial muscle activity emerges with full contribution of these systems so that the effects of reflex delays become negligible. We tested the hypothesis that threshold control of muscle activation may be instrumental in preventing destabilizing effects of proprioceptive delays in spinal and trans-cortical pathways to motoneurons. The analysis was made by recording fast elbow movements (peak velocity approximately 300-500 degrees/s) and simulating them in a dynamic model that incorporates the notion of threshold control of intrinsic and reflex muscle properties. The model was robust in reproducing experimental movement patterns (R (2)>0.95). It generated stable output despite substantial proprioceptive (up to 100 ms) and electromechanical (40 ms) delays. Stability was thus ensured for delays not only in segmental (about 25-50 ms) but also in trans-cortical loops (50-70 ms). Our study illustrates that a natural physiological process--threshold control--may manifest feed-forward properties hitherto attributed to hypothetical internal neural models.

It has been suggested that multijoint movements result from the specification of a referent configuration of the body. The activity of muscles and forces required for movements emerge depending on the difference between the actual and referent body configurations. We identified the referent arm configurations specified by the nervous system to bring the arm to the target position both in healthy individuals and in those with arm motor paresis due to stroke. From an initial position of the right arm, subjects matched a force equivalent to 30% of their maximal voluntary force in that position. The external force, produced at the handle of a double-joint manipulandum by two torque motors, pulled the hand to the left (165 degrees ) or pushed it to the right (0 degrees ). For both the initial conditions, three directions of the final force (0 degrees , +20 degrees , and -20 degrees ) with respect to the direction of the initial force were used. Subjects were instructed not to intervene when the load was unexpectedly partially or completely removed. Both groups of subjects produced similar responses to unloading of the double-joint arm system. Partial removal of the load resulted in distinct final hand positions associated with unique shoulder-elbow configurations and joint torques. The net static torque at each joint before and after unloading was represented as a function of the two joint angles describing a planar surface or invariant characteristic in 3D torque/angle coordinates. For each initial condition, the referent arm configuration was identified as the combination of elbow and shoulder angles at which the net torques at the two joints were zero. These configurations were different for different initial conditions. The identification of the referent configuration was possible for all healthy participants and for most individuals with hemiparesis suggesting that they preserved the ability to adapt their central commands-the referent arm configurations-to accommodate changes in external load conditions. Despite the preservation of the basic response patterns, individuals with stroke damage had a more restricted range of hand trajectories following unloading, an increased instability around the final endpoint position, altered patterns of elbow and shoulder muscle coactivation, and differences in the dispersion of referent configurations in elbow-shoulder joint space compared to healthy individuals. Moreover, 4 out of 12 individuals with hemiparesis were unable to specify referent configurations of the arm in a consistent way. It is suggested that problems in the specification of the referent configuration may be responsible for the inability of some individuals with stroke to produce coordinated multijoint movements. The present work adds three findings to the motor control literature concerning stroke: non-significant torque/angle relationships in some subjects, narrower range of referent arm configurations, and instability about the final position. This is the first demonstration of the feasibility of the concept of the referent configuration for the double-joint muscle-reflex system and the ability of some individuals with stroke to produce task-specific adjustments of this configuration.

In addition to local biomechanical and reflex factors influencing muscle activation, global factors may be used by the nervous system to control all muscles in a coherent and task-specific way. It has been hypothesized that a virtual or referent (R) configuration of the body determined by muscle recruitment thresholds specified by neural control levels is such a factor. Due to the threshold nature of the R configuration, the activity of each muscle depends on the difference between the actual (Q) and the R configuration of the body. The nervous system modifies the R configuration to produce movement. One prediction of this hypothesis is that the Q and R configurations may match each other, most likely in movements with reversals in direction, resulting in a minimum in the electromyographic (EMG) activity level of muscles involved. The depth of the minima is constrained by the degree of coactivation of opposing muscle groups. Another prediction is that EMG minima in the activity of multiple muscles may occur not only when the movement is assisted but also when it is opposed by external forces (e.g., gravity). To verify these predictions, we analyzed EMG patterns of 16-21 functionally diverse muscles of the legs, trunk, and arms during jumping and stepping in place. One EMG minimum in the activity of all muscles regularly occurred near the apex of the jump. A minimum was also observed near the point of transition of the body from flexion to extension leading to a jump. During stepping in place, the activity of muscles of each side of the body was usually minimized near the beginning and near the end of the stance phase as well as during the maximum elevation of the foot. Since EMG minima occurred not only during gravity-assisted but also gravity-opposed movement reversals, it is concluded that neural factors (such as matching between the Q and R) rather than mechanical factors are responsible for minimizing the EMG activity in these movements.

The role of the basal ganglia in the adaptive control of movement was investigated by unexpectedly perturbing movements in 8 patients with Parkinson's disease (PD) tested off medication and in 6 aged-matched healthy subjects. Subjects performed two movement components simultaneously and without visual feedback: touching the nose with the finger while leaning the trunk forward. Subjects wore a harness connected to an electromagnet, which was attached to a wall. The trunk movement was mechanically blocked in randomly selected trials by engaging the electromagnet. While healthy subjects performed the task equally well in both conditions, PD subjects' hand movements significantly deteriorated in trunk-perturbed compared to trunk-free trials. Deteriorated hand movements were characterized by segmented hand paths, unsmooth velocity profiles, and prolonged movement times. This finding indicated that the relatively local trunk perturbation had a global effect on the hand movement of PD subjects, necessitating them to reinitiate, after some delay, their arm movement in perturbed trials. Thus, the basal ganglia may be a critical node in brain networks mediating the flexibility of responses to altered motor states.

During trunk-assisted reaching to targets placed within arm's length, the influence of trunk motion on the hand trajectory is compensated for by changes in the arm configuration. The role of proprioception in this compensation was investigated by analyzing the movements of 2 deafferented and 12 healthy subjects. Subjects reached to remembered targets (placed approximately 80 degrees ipsilateral or approximately 45 degrees contralateral to the sagittal midline) with an active forward movement of the trunk produced by hip flexion. In 40% of randomly selected trials, trunk motion was mechanically blocked. No visual feedback was provided during the experiment. The hand trajectory and velocity profiles of healthy subjects remained invariant whether or not the trunk was blocked. The invariance was achieved by changes in arm interjoint coordination that, for reaches toward the ipsilateral target, started as early as 50 ms after the perturbation. Both deafferented subjects exhibited considerable, though incomplete, compensation for the effects of the perturbation. Compensation was more successful for reaches to the ipsilateral target. Both deafferented subjects showed invariance between conditions (unobstructed or blocked trunk motion) in their hand paths to the ipsilateral target, and one did to the contralateral target. For the other deafferented subject, hand paths in the two types of trials began to deviate after about 50% into the movement, because of excessive elbow extension. In movements to the ipsilateral target, when deafferented subjects compensated successfully, the changes in arm joint angles were initiated as early as 50 ms after the trunk perturbation, similar to healthy subjects. Although the deafferented subjects showed less than ideal compensatory control, they compensated to a remarkably large extent given their complete loss of proprioception. The presence of partial compensation in the absence of vision and proprioception points to the likelihood that not only proprioception but also vestibulospinal pathways help mediate this compensation.

International audience; Recent studies have shown that the hand-pointing movements within arm's reach remain invariant whether the trunk is recruited or not or its motion is unexpectedly prevented. This suggests the presence of compensatory arm-trunk coordination minimizing the deflections of the hand from the intended trajectory. It has been postulated that vestibular signals elicited by the trunk motion and transmitted to the arm motor system play a major role in the compensation. One prediction of this hypothesis is that vestibular stimulation should influence arm posture and movement during reaching. It has been demonstrated that galvanic vestibular stimulation (GVS) can influence the direction of pointing movements when body motion is restrained. In the present study, we analyzed the effects of GVS on trunk-assisted pointing movements. Subjects either moved the hand to a target or maintained a steady-state posture near the target, while moving the trunk forward with the eyes closed. When GVS was applied, the final position of the hand was deviated in the lateral and sagittal direction in both tasks. This was the result of two independent effects: a deviation of the trunk trajectory and a modification of the arm position relative to the trunk. Thus, the vestibular system might be directly involved not only in the control of trunk motion but also in the arm-trunk coordination during trunk-assisted reaching movements.

Redundancy is associated with the ability of the nervous system to select different interjoint coordinations and movement trajectories to achieve the same motor goal. The nervous system may coordinate multiple degrees of freedom (DF) by combining them in a task-specific way to control them as a unit or synergy. Some movements may be accomplished using only one synergy, whereas other movements may employ several synergies. To investigate the problem of interjoint coordination, we applied principal component (PC) analysis to eight types of movement in healthy male subjects: forward squats, backward squats, sideways squats, squats on one leg, walking three steps, stepping in place, going up a step, and going down a step. Angular changes in four DF were analyzed: thigh flexion-extension, knee flexion-extension, ankle flexion-extension, thigh abduction-adduction, with the former three DF investigated in all movements. For many movements, two synergies were sufficient to account for more than 95% of DF angular excursions. Squatting on one leg could be described using only one synergy (99%). The angle between the vectors representing PCs for movements produced with the right and left legs could be less than 10 degrees for some movements but could reach 25 degrees for other movements. The nervous system may thus use somewhat different interjoint coordinations while producing movements on the right and the left sides. The angle between the first PCs of different movements could be smaller than 10 degrees. Thus there may be a common but adjustable basic synergy that is used to produce different movements. Additional synergies provide the transition from one movement to another.

Peripheral (muscle) aspects of fatigue are well documented. However, little is known about the central aspects of fatigue that could influence, in particular, multijoint coordination. To investigate the central aspects of fatigue, we compared the multijoint kinematics of non-fatigued and fatigued individuals while sawing. Muscle fatigue was associated with decreases in sawing force and movement amplitude at the elbow whereas the basic characteristics of the saw trajectory, including the movement direction, extent and duration, remained invariant. This invariance was maintained by increasing the movement amplitude at the wrist, shoulder and trunk. The system thus takes advantage of the redundancy of the motor apparatus to maintain the endpoint trajectory despite fatigue.

According to previous studies, muscles become active in response to deviations from a threshold (referent) position of body segments. To test the hypothesis that corticospinal pathways set and reset the referent position in a task-specific way, we evaluated corticospinal influences at wrist positions established before and after voluntary motion as well as before and after involuntary motion elicited by the sudden removal of a load. Although tonic electromyogram (EMG) levels at pre- and post-unloading wrist positions were different, the corticospinal influences and thus the referent wrist position remained the same. These influences and the referent position changed, however, when subjects voluntarily moved their wrists to the other position. Thus, referent control strategies underlying the two types of motor actions are fundamentally different. We also tested the hypothesis that somatosensory afferents inform the brain about the deviation (P) of body segments from the centrally set referent position, R. To identify the actual position (Q) of body segments and form the position sense (PS), the central and afferent signals are combined. The PS rule was confirmed by demonstrating that: subjects are able to successfully reproduce involuntary changes in position elicited by sudden unloading of wrist flexors by making voluntary wrist movements; successfully reproduce elbow joint positions under different constant loads; subjects may not be aware of elbow flexion elicited by tendon vibration until a certain limit (no-motion illusion). In conclusion, R and P are additive components of PS and, contrary to conventional assumptions, PS is independent of sense of effort or efference copy.

Movements of different body segments may be combined in different ways to achieve the same motor goal. How this is accomplished by the nervous system was investigated by having subjects make fast pointing movements with the arm in combination with a forward bending of the trunk that was unexpectedly blocked in some trials. Subjects moved their hand above the surface of a table without vision from an initial position near the midline of the chest to remembered targets placed within the reach of the arm in either the ipsi- or contralateral workspace. In experiment 1, subjects were instructed to make fast arm movements to the target without corrections whether or not the trunk was arrested. Only minor changes were found in the hand trajectory and velocity profile in response to the trunk arrest, and these changes were seen only late in the movement. In contrast, the patterns of the interjoint coordination substantially changed in response to the trunk arrest, suggesting the presence of compensatory arm-trunk coordination minimizing the deflections from the hand trajectory regardless of whether the trunk is recruited or mechanically blocked. Changes in the arm interjoint coordination in response to the trunk arrest could be detected kinematically at a minimal latency of 50 ms. This finding suggests a rapid reflex compensatory mechanism driven by vestibular and/or proprioceptive afferent signals. In experiment 2, subjects were required, as soon as they perceived the trunk arrest, to change the hand motion to the same direction as that of the trunk. Under this instruction, subjects were able to initiate corrections only after the hand approached or reached the final position. Thus, centrally mediated compensatory corrections triggered in response to the trunk arrest were likely to occur too late to maintain the observed invariant hand trajectory in experiment 1. In experiment 3, subjects produced similar pointing movements, but to a target that moved together with the trunk. In these body-oriented pointing movements, the hand trajectories from trials in which the trunk was moving or arrested were substantially different. The same trajectories represented in a relative frame of reference moving with the trunk were virtually identical. We conclude that hand trajectory invariance can be produced in an external spatial (experiment 1) or an internal trunk-centered (experiment 3) frame of reference. The invariance in the external frame of reference is accomplished by active compensatory changes in the arm joint angles nullifying the influence of the trunk motion on the hand trajectory. We suggest that to make a transition to the internal frame of reference, control systems suppress this compensation. One of the hypotheses opened to further experimental testing is that the integration of additional (trunk) degrees of freedom into movement is based on afferent (proprioceptive, vestibular) signals stemming from the trunk motion and transmitted to the arm muscles.

When arm and trunk segments are involved in reaching for objects within arm's reach, vestibulospinal pathways compensate for trunk motion influence on arm movement. This compensatory arm-trunk synergy is characterised by a gain coefficient of 0 to 1. Vestibular patients have less efficient arm-trunk synergies and lower gains. To assess the clinical usefulness of the gain measure, we used a portable ultrasound-based device to characterize arm-trunk coordination deficits in vestibular patients. Arm-trunk coordination without vision was measured in a Stationary Hand Task where hand position was maintained during trunk movement, and a Reaching Task with and without trunk motion. Twenty unilateral vestibular patients and 16 controls participated. For the Stationary Hand task, patient gains ranged from g=0.94 (good compensation) to 0.31 (poor compensation) and, on average, were lower than in controls (patients: 0.67 ± 0.19; controls: 0.85 ± 0.07; p< 0.01). Gains were significantly correlated with clinical tests (Sensory Organization; r=0.62, p< 0.01, Foam Romberg Eyes Closed; r=0.65, p< 0.01). For the Reaching Task, blocking trunk movement during reaching modified hand position significantly more in patients (8.2 ± 4.3 cm) compared to controls (4.5 ± 1.7 cm, p< 0.02) and the amount of hand position deviation was correlated with the degree of vestibular loss in a sub-group (n=14) of patients. Measurement of the Stationary Task arm-trunk gain and hand deviations during the Reaching Task can help characterize sensorimotor problems in vestibular-deficient patients and track recovery following therapeutic interventions. The ultrasound-based portable device is suitable for measuring vestibulospinal deficits in arm-trunk coordination in a clinical setting.

Previous studies addressing the problem of the control of multiple degrees of freedom have examined the influence of trunk movement on pointing movements within the arm's reach. Such movements may be controlled by two functionally independent units of coordination (synergies): one involving only arm joints and producing the hand trajectory to the target (the transport synergy), and the other coordinating trunk and arm movements leaving the hand trajectory unchanged (the compensatory synergy). The question of whether or not this functional subdivision depends on visual feedback was addressed in the present study. We also tested whether or not the motor effects of different synergies are summated as independent components, a control strategy called "superposition." Finally, we investigated whether or not the relationship between different degrees of freedom within each synergy could be considered linear resulting in proportional changes in different joint angles. Seated subjects produced fast, uncorrected arm movements to an ipsi- or a contralateral target in the direction of +/-45 degrees to the sagittal midline of the trunk. Targets could be reached using the arm alone (control trials) or by combining the arm motion with a forward or backward trunk motion produced by hip flexion or extension (test trials), with and without visual feedback. The shape of the hand trajectory, its direction and tangential velocity, movement precision, joint angles and the sequence of the trunk and hand recruitment and de-recruitment were measured. In both visual conditions, the direction of the hand trajectory observed in control trials was generally preserved in test trials. In terms of sequencing, even in the absence of vision, the trunk movement was initiated before the onset of and outlasted the hand shift, indicating that the potential influence of the trunk on the hand movement was compensated by rotations in the elbow and shoulder joint. The analysis of other variables also implied that the effects of trunk recruitment on the hand trajectory were minor compared to those which could be observed if these effects were not compensated by appropriate changes in the arm joint angles. It was concluded that an arm-trunk compensatory synergy is present in pointing movements regardless of visual feedback. Principal component analysis showed that the relationship between elbow, shoulder and hip joint angles in individual arm and combined arm-trunk movements cannot be considered linear, implying that this relationship is adjusted according to the changing arm geometry. The changes in each arm joint angle (elbow, shoulder) elicited by a forward trunk bending in one block of trials were compared with those elicited by a backward bending in another block, whereas the hand moved to the same target in both blocks. These changes were opposite but of similar magnitude. As a result, for each moment of movement, the mean joint angle obtained by averaging across two directions of trunk motion was practically identical to that in control trials in which the trunk was motionless. It is concluded that the transport and arm-trunk compensatory synergies are combined as independent units, according to the principle of superposition. This principle may simplify the control of the coordination of a redundant number of degrees of freedom.

It has been suggested that the coordination of the activity of multiple muscles results from the comparison of the actual configuration of the body with a referent configuration specified by the nervous system so that the recruitment and gradation of the activity of each skeletal muscle depend on the difference between these two configurations. Active movements may be produced by the modification of the referent configuration. The hypothesis predicts the existence of a global minimum in electromyographic (EMG) activity of multiple muscles during movements involving reversals in direction. This prediction was tested in five subjects by analysing movements resembling the act of reaching for an object placed beyond one's reach from a sitting position. In such movements, initially sitting subjects raise their body to a semi-standing position and then return to sitting. Consistent with the hypothesis is the observation of a global minimum in the surface EMG activity of 16 muscles of the arm, trunk and leg at a specific phase of the movement. When the minimum occurred, EMG activity of each muscle did not exceed 2–7% of its maximal activity during the movement. As predicted, global EMG minima occurred at the phase corresponding to the reversal in movement direction, that is, during the transition from raising to lowering of the body. The global EMG minimum may represent the point at which temporal matching occurs between the actual and the referent body configurations. This study implies a specific link between motor behavior and the geometric shape of the body modified by the brain according to the desired action.

This study examined the influence of trunk recruitment on the kinematic characteristics of pointing movements. The distribution of final positions of the hand, the extent and direction of the hand trajectory was basically preserved when trunk movement was combined with arm pointing. These effects were observed during pointing not only with but also without vision. The results imply that two functionally independent units of coordination are used in pointing regardless of visual feedback-one producing arm movement to the target (the reaching synergy) and the other coordinating trunk and arm movements diminishing the influence of the trunk on the arm endpoint trajectory (the compensatory synergy).

We investigated phasic and tonic stretch reflexes in human jaw-opener muscles, which have few, if any, muscle spindles. Jaw-unloading reflexes were recorded for both opener and closer muscles. Surface electromyographic (EMG) activity was obtained from left and right digastric and superficial masseter muscles, and jaw orientation and torques were recorded. Unloading of jaw-opener muscles elicited a short-latency decrease in EMG activity (averaging 20 ms) followed by a short-duration silent period in these muscles and sometimes a short burst of activity in their antagonists. Similar behavior in response to unloading was observed for spindle-rich jaw-closer muscles, although the latency of the silent period was statistically shorter than that observed for jaw-opener muscles (averaging 13 ms). Control studies suggest that the jaw-opener reflex was not due to inputs from either cutaneous or periodontal mechanoreceptors. In the unloading response of the jaw openers, the tonic level of EMG activity observed after transition to the new jaw orientation was monotonically related to the residual torque and orientation. This is consistent with the idea that the tonic stretch reflex might mediate the change in muscle activation. In addition, the values of the static net joint torque and jaw orientation after the dynamic phase of unloading were related by a monotonic function resembling the invariant characteristic recorded in human limb joints. The torque-angle characteristics associated with different initial jaw orientations were similar in shape but spatially shifted, consistent with the idea that voluntary changes in jaw orientation might be associated with a change in a single parameter, which might be identified as the threshold of the tonic stretch reflex. It is suggested that functionally significant phasic and tonic stretch reflexes might not be mediated exclusively by muscle spindle afferents. Thus, the hypothesis that central modifications in the threshold of the tonic stretch reflex underlie the control of movement may be applied to the jaw system.

Using a non-linear dynamic model based on the lambda version of the equilibrium-point hypothesis, we investigated the shape and duration of the control patterns underlying discrete elbow movements. The model incorporates neural control variables, time-, position- and velocity-dependent intrinsic muscle and reflex properties. Two control variables (R and C) specify a positional frame of reference for activation of flexor and extensor motoneurons. The variable R (reciprocal command) specifies the referent joint angle (R) at which the transition of net flexor to extensor active torque or vice versa can be observed during changes in the actual joint angle elicited by an external force. The variable C (coactivation command) surrounds the transition angle by an angular range in which flexor and extensor muscles may be simultaneously active (if C0) or silent (if Cor = 0). An additional, time-dimensional control variable (mu command) influences the dependency of the threshold of the stretch reflex on movement velocity. This control variable is responsible for the reflex damping. Changes in the R command result in shifts in the equilibrium state of the system, a dynamical process leading to electromyographic modifications and movement production. Commands C and mu provide movement stability and effective energy dissipation preventing oscillations at the end of movement. A comparison of empirical and model data was carried out. A monotonic ramp-shaped pattern of the R command can account for the empirical kinematic and electromyographic patterns of the fastest elbow flexion movements made with or without additional inertia, as well as of self-paced movements. The rate of the shifts used in simulation was different for the three types of movements but independent of movement distance (20-80 degrees). This implies that, for a given type of movement, the distance is encoded by the duration of shift in the equilibrium state. The model also reproduces the kinematic and electromyographic patterns of the fastest uncorrected movements opposed in random trials by a high load (80-90% of the maximal) generated by position feedback to a torque motor. The following perturbation effects were simulated: a substantial decrease in the arm displacement (from 60-70 degrees to 5-15 degrees) and movement duration (to about 100 ms) so that these movements ended near the peak velocity of those which were not perturbed; a prolongation of the first agonist electromyographic burst as long as the load was applied; the suppression of the antagonist burst during the dynamic and static phases of movements: the reappearance of the antagonist burst in response to unloading accompanied by a short-latency suppression of agonist activity. These kinematic and electromyographic features of both perturbed and non-perturbed movements were reproduced by using the same control patterns which elicited a monotonic shift in the equilibrium state of the system ending before the peak velocity of non-perturbed movements. Our results suggest that the neural control processes underlying the fastest unopposed changes in the arm position are completed long before the end of the movement and phasic electromyographic activity. Neither the timing nor the amplitude of electromyographic bursts are planned but rather they represent the long-lasting dynamic response of central, reflex and mechanical components of the system to a monotonic, short-duration shift in the system's equilibrium state.

Abnormalities of muscle tone are an integral component of many chronic motor disorders affecting the central nervous system (CNS) in children and adults. Excessive, disabling muscle tone is called ‘spasticity’. Spasticity can interfere with movement and can lead to stiff, painful joints [1]. Over half a million people in the United States and 12 million worldwide are affected by spasticity. Spasticity results from dysgenesis or injury to sensorimotor pathways in the cortex, basal ganglia, thalamus, cerebellum, brainstem, central white matter, or spinal cord. Injury occurring in children is known as cerebral palsy (CP) [2]. In adults, CNS injury or disease associated with spasticity includes stroke, spinal cord injury, multiple sclerosis and traumatic brain injury. Although the presence or absence of spasticity can be identified using current clinical scales, the accuracy of determination of spasticity severity and the relationship between severity level and deficits of voluntary movements remain elusive (see [3,4]). These questions are of interest to both researchers and clinicians, because the precise quantification of spasticity is important to establish medical and physical therapeutic effectiveness. We designed a new measure of spasticity, called the Montreal Stretch Reflex Threshold (MSRT) measure, based on a) Lance’s definition [5] of spasticity as "a motor disorder characterized by a velocity-dependent increase in tonic stretch reflexes with exaggerated tendon jerks, resulting from hyperexcitability of the stretch reflex arc" and b) the threshold control theory of motor control [6]. This concept of measurement based on the velocitydependent stretch reflex threshold is a departure from the more common approach that characterizes the resistance to stretch of the passive muscle (e.g., [7]) and is likely to be more related to motor control deficits than resistance measures. If so, this may lead to a better ability to diagnose the motor impairment in spasticity and may lead to improved patient care. Thus, instead of determining the level of the resistance to stretch of the muscle as is usually done to assess spasticity, the main measure in the new device is the threshold joint angle at which the tonic stretch reflex (TSR) starts to produce resistance, which is clinically identified as spasticity.

According to recent findings, intentional motor actions are controlled by resetting the referent position, R, at which neuromuscular elements, including reflexes, begin to act. It is suggested that somatosensory afferents inform the brain about the deviation (P) of body segments from the centrally set referent position. To perceive the actual position (Q) of body segments and form the position sense (PS), the central and afferent signals are combined: Q=R+P. In previous studies, the R has been shown to remain invariant during involuntary changes in the wrist position elicited by sudden unloading of muscles, suggesting that only the afferent component is responsible for the PS during this reflex. In contrast, the central PS component, R, is predominantly responsible for PS during intentional motion in isotonic conditions. We tested the hypothesis that the R and P are interchangeable PS components such that involuntary changes in wrist position elicited by the unloading reflex can easily be reproduced by making intentional changes in wrist position in isotonic conditions, in the absence of vision. The PS rule also suggests that PS is independent of sense of effort, which was tested by asking subjects to reproduce elbow joint angles under different constant loads. We also tested the hypothesis that tendon vibration may elicit motion that may not be perceived by subjects (no-motion illusion). These hypotheses were confirmed in three experiments. It is concluded that the R and P are additive components of PS and that, contrary to the conventional view, PS is independent of the sense of effort or efference copy. The PS rule also explains kinesthetic illusions and the phantom limb phenomenon. This study advances the understanding of action–perception coupling in kinesthesia.

The cerebellum probably obeys the rules of sensorimotor integration common in the nervous system. One such a rule is formulated: the nervous system organizes spatial frames of reference for the sensorimotor apparatus and produces voluntary movements by shifting their origin points. We give examples of spatial frames of reference for different single- and multi-joint movements including locomotion and also illustrate that the process of motor development and learning may depend critically on the formation of appropriate frames of reference and the organism's ability to manage them. We suggest that a solution to the problem of sensorimotor integration may not be trivial and may actually change the mental and experimental paradigms used in the understanding of the cerebellum and other brain structures, [HOUK et al.; SIMPSON et al.; SMITH; TIIACH]

Sensorimotor mechanisms were studied on the basis of kinematic and electromyographic data as well as the static torque developed by the muscles as a function of joint angle. The latter relationship is known as the torque/angle characteristic. Fast single-joint movement may result from a shift in this characteristic and a change in its slope. Such movements were studied at the wrist in 9 normal and 1 deafferented subject. After training to flex the wrist to a target, subjects repeated the same movements but in random test trials movements were opposed by the load generated by linear position feedback to a torque motor. At the end of the loaded trials, the load was suddenly removed. In the second experiment, subjects made wrist movements to the target that were opposed by the load and, on random test trials, the movements were not loaded. In these test trials, the wrist arrived in a static position outside the target zone. In both experiments, subjects were instructed not to correct errors. The final torque/angle characteristics specified in the movements were reconstructed on the basis of the static wrist positions and torques before and after unloading. Normal subjects made movements by shifting the position of the torque/angle characteristic and by increasing its slope. If subjects indeed maintained the same pattern of control variables (descending commands), the same final position of the characteristic would be reproduced from trial to trial regardless of load perturbations. This assumption of equifinality was tested by comparing the final position of the wrist in nonloaded movements with that after removal of the load in loaded movements. Equifinality was observed in normal subjects. Movements in the deafferented subject were also associated with a shift of the torque/angle characteristic and a change in its slope. However, she was unable to consistently reproduce its final position. In spite of muscle coactivation, her maximal stiffness was lower than in normal subjects. In the absence of vision, the subject made movements with the load by increasing the slope of the characteristic instead of by shifting its position far enough. Load perturbation affected her final wrist position (inequifinality), which may reflect the presence of a significant hysteresis of the characteristic as a result of the absence of stretch reflexes. The deficits following deafferentation presumably result from the destruction of biomechanical and sensorimotor mechanisms including the ability of control variables to specify the positional frame of reference for afferent and descending systems.Key words: sensorimotor integration, positional frames of reference, deafferentation, wrist stiffness, equilibrium-point hypothesis.

Some hypotheses suggest that stretch reflex threshold regulation may be an essential element of motor control. Disturbances in this mechanism may lead to motor dysfunction. We investigated this possibility by comparing stretch reflex threshold regulation in 11 spastic hemiparetic and 6 normal subjects. Subjects sat with their arms fully supported in a forearm and hand mold attached to a manipulandum mounted on and controlled by a torque motor. They remained completely passive while their elbow was extended from 30 degrees flexion through an arc of 100 degrees. Displacement and velocity of the forearm were measured as well as EMG signals from 2 elbow flexors and 2 elbow extensors, when the elbow flexors were stretched at each of 7 velocities. Velocities ranged from 8 to 160 degrees/s for hemiparetic subjects and from 32 to 300 degrees/s for normal subjects. Phase diagrams (velocity versus angle) were plotted and the threshold angles (lambda) for muscle activation at each velocity of stretch were used to determine the static stretch reflex threshold (lambda) and the slope (mu) of the relationship between the lambda s and velocity. Our main findings were that static and dynamic stretch reflex thresholds were decreased in spastic hemiparetic compared to normal subjects and that the thresholds depended on velocity. The static threshold value correlated with the severity of clinically measured spasticity. In addition, the range of regulation of lambda was decreased in the patients compared to normal. This may explain some of the problems of force and position regulation as well as hypertonus (and weakness) common to these patients.

Previous findings suggest that, by influencing the subthreshold state of motoneurons, the corticospinal pathways can set and reset the threshold position at which wrist muscle recruitment begins. Here we assumed that the corticospinal system can change the threshold position in a similar way before anticipated perturbation to pre-determine an appropriate emerging response to it. We first analyzed motor-evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS) applied to the wrist area of motor cortex before unloading of preloaded wrist flexors, i.e. before the subsequent involuntary wrist motion to another position (natural unloading). Subjects then learned to diminish the post-unloading movement extent without activating antagonist (extensor) muscles before unloading or making intentional movement corrections after unloading (adjusted unloading). Although activity levels of wrist muscles before unloading were similar, MEPs of extensor but not pre-loaded flexor muscles were higher before adjusted unloading. We also applied TMS in combination with a torque pulse that shortened extensor muscles such that the MEP occurred when the motoneuronal excitability was minimized. Although diminished following muscle shortening, MEPs before adjusted unloading were still higher than before natural unloading. Results suggest that the corticospinal system, possibly together with other descending systems participated in the tonic subthreshold facilitation of antagonist motoneurons before adjusted unloading, which appears sufficient in modifying motor commands and motion leading to adjusted unloading. This study reinforces previous findings that descending systems, in particular, the corticospinal system can employ threshold position control during and after learning a novel action.