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The referent control theory (RCT) for action and perception is an advanced formulation of the equilibrium-point hypothesis. The RCT suggests that rather than directly specifying the desired motor outcome, the nervous system controls action and perception indirectly by setting the values of parameters of physical and physiological laws. This is done independently of values of kinematic and kinetic variables including electromyographic patterns describing the motor outcome. One such parameter—the threshold muscle length, λ, at which motoneurons of a given muscle begin to be recruited, has been identified experimentally. In RCT, a similar parameter, the referent arm position, R, has been defined for multiple arm muscles as the threshold arm position at which arm muscles can be quiescent but activated depending on the deflection of the actual arm position, Q, from R. Changes in R result in reciprocal changes in the activity of opposing muscle groups. We advanced the explanatory power of RCT by combining the usual biomechanical descriptions of motor actions with the identification of the timing of R underlying arm movements made with reversals in three directions and to three different extents. We found that in all movements, periods of minimization of the activity of multiple muscles could be identified at ∼61%–86% of the reaching extent in each direction. These electromyographic minimization periods reflect the spatial coordinates at which the R and Q overlap during the production of movements with reversals. The findings support the concept of the production of arm movement by shifting R.

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.

We tested the hypothesis that the ipsilateral corticospinal system, like the contralateral corticospinal system, controls the threshold muscle length at which wrist muscles and the stretch reflex begin to act during holding tasks. Transcranial magnetic stimulation was applied over the right primary motor cortex in 21 healthy subjects holding a smooth or coarse block between the hands. Regardless of the lifting force, motor evoked potentials in right wrist flexors were larger for the smooth block. This result was explained based on experimental evidence that motor actions are controlled by shifting spatial stretch reflex thresholds. Thus, the ipsilateral corticospinal system is involved in threshold position control by modulating facilitatory influences of hand skin afferents on motoneurons of wrist muscles during bimanual object manipulation.

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.

Unperturbed human locomotion presumably results from feedforward shifts in stable body equilibrium in the environment, thus avoiding falling and subsequent catching considered in alternative theories of locomotion. Such shifts are achieved by relocation of the referent body configuration at which multiple muscle recruitment begins. Rather than being directly specified by a central pattern generator, multiple muscles are activated depending on the extent to which the body is deflected from the referent, threshold body configuration, as confirmed in previous studies. Based on the referent control theory of action and perception, solutions to classical problems in motor control are offered, including the previously unresolved problem of the integration of central and reflex influences on motoneurons and the problem of how posture and movement are related. The speed of locomotion depends on the rate of shifts in the referent body configuration. The transition from walking to running results from increasing the rate of referent shifts. It is emphasised that there is a certain hierarchy between reciprocal and co-activation of agonist and antagonist muscles during locomotion and other motor actions, which is also essential for the understanding of how locomotor speed is regulated. The analysis opens a new avenue in neurophysiological approaches to human locomotion with clinical implications.

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

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.

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.

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.

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.

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.

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.

Advances to the equilibrium-point (EP) theory and solutions to several classical problems of action and perception are suggested and discussed. Among them are (1) the posture-movement problem of how movements away from a stable posture can be made without evoking resistance of posture-stabilizing mechanisms resulting from intrinsic muscle and reflex properties; (2) the problem of kinesthesia or why our sense of limb position is fairly accurate despite ambiguous positional information delivered by proprioceptive and cutaneous signals; (3) the redundancy problems in the control of multiple muscles and degrees of freedom. Central to the EP hypothesis is the notion that there are specific neural structures that represent spatial frames of reference (FRs) selected by the brain in a task-specific way from a set of available FRs. The brain is also able to translate or/and rotate the selected FRs by modifying their major attributes-the origin, metrics, and orientation-and thus substantially influence, in a feed-forward manner, action and perception. The brain does not directly solve redundancy problems: it only limits the amount of redundancy by predetermining where, in spatial coordinates, a task-specific action should emerge and allows all motor elements, including the environment, to interact to deliver a unique action, thus solving the redundancy problem (natural selection of action). The EP theory predicts the existence of specific neurons associated with the control of different attributes of FRs and explains the role of mirror neurons in the inferior frontal gyrus and place cells in the hippocampus. WIREs Cogni Sci 2011 2 287-304 DOI: 10.1002/wcs.108 For further resources related to this article, please visit the WIREs website.

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.

According to a view that has dominated the field for over a century, the brain programs muscle commands and uses a copy of these commands [efference copy (EC)] to adjust not only resulting motor action but also ongoing perception. This view was helpful in formulating several classical problems of action and perception: (1) the posture-movement problem of how movements away from a stable posture can be made without evoking resistance of posture-stabilizing mechanisms resulting from intrinsic muscle and reflex properties; (2) the problem of kinesthesia or why our sense of limb position is good despite ambiguous positional information delivered by proprioceptive and cutaneous signals; (3) the problem of visual space constancy or why the world is perceived as stable while its retinal image shifts following changes in gaze. On closer inspection, the EC theory actually does not solve these problems in a physiologically feasible way. Here solutions to these problems are proposed based on the advanced formulation of the equilibrium-point hypothesis that suggests that action and perception are accomplished in a common spatial frame of reference selected by the brain from a set of available frames. Experimental data suggest that the brain is also able to translate or/and rotate the selected frame of reference by modifying its major attributes-the origin, metrics and orientation-and thus substantially influence action and perception. Because of this ability, such frames are called physical to distinguish them from symbolic or mathematical frames that are used to describe system behavior without influencing this behavior. Experimental data also imply that once a frame of reference is chosen, its attributes are modified in a feedforward way, thus enabling the brain to act in an anticipatory and predictive manner. This approach is extended to sense of effort, kinesthetic illusions, phantom limb and phantom body phenomena. It also addresses the question of why retinal images of objects are sensed as objects located in the external, physical world, rather than in internal representations of the brain.

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.

This quotation (http://www.quotationspage.com/faq.php#85) is reminiscent of the Godel’s incompleteness theorem in mathematics: In any axiomatic system of postulates there would always be some statements the validity of which could not be established within this system (Kennedy 2011). Einstein’s quote points to similar limitations in other branches of science and everyday life.

Some ideas outlined in this book emerged in the unique scientific atmosphere created by a group of scientists that is now known in the West as the Moscow Biological School. As an historical and scientific phenomenon, this School deserves to be described in a separate book (Berkinblit and Latash 2005; see also Stuart 2005). As a proud member of this School, I will briefly describe it to acknowledge its influence on my thinking and to express my gratitude to its members. I will also describe the circumstances related to my decision to switch from physics to neuroscience. This Chapter is not entirely biographical: I introduce some notions related to the idea of referent control described in this book.

The co-existence of divergent theories of action and perception is indicative of a confusion, rather than of a consensus in the understanding of basic principles of brain functioning. This situation in behavioral neurosciences is comparable to one that existed in the natural sciences before Galileo and Newton. This book is an attempt to overcome this situation by formulating physiologically feasible principles underlying action and perception. I take advantage of my background in physics and neurophysiology (see Chap. 1) to approach this challenging task and formulate several criteria for accepting/rejecting diverse theories of action and perception. One criterion is well-known: Theories should pass the scrutiny of the scientific method established by Galileo. Another, less known criterion is based on the recognition that physical laws (also called natural or universal laws) constrain the ways in which biological systems can accomplish action and perception. Finally, it is taken for granted that any theory of action and perception should be physiologically feasible. These criteria provide a natural framework to be respected in evaluating existing and creating new theories of action and perception.

In this book, I have described a theory of action and perception—referent control. Strictly based on experimental material, the theory places action and perception in the context of physical laws to offer solutions of several classical problems in behavioral neuroscience. This approach also results in the rejection of theories in behavioral neuroscience that are rooted in mechanical reductionism associated with the idea that neural control levels are directly involved in pre-programming of mechanical variables and patterns of motor commands to muscles required for reaching the motor goal. In contrast, these levels only pre-determine the spatial frame of reference and its attributes such as its origin, orientation and metrics to indicate where, in this frame, a motor action can emerge due to natural interactions of the organism with the environment. If necessary, action corrections are also accomplished indirectly, at a referent level.

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.

A specific form of parametric control described in the previous section—referent or threshold position control of motor actions—can be considered as the foundation of a physiologically feasible theory of action. As a first step towards this goal, we need to explain how electro-chemical mono- and poly-synaptic influences on α-MNs from descending and spinal centers are transformed into spatial variables ˗ threshold angle, R, or threshold muscle length, λ. The referent C command is also a spatial variable: In the case of simultaneous facilitation of agonist and antagonist MNs, it subdivides the biomechanical range into three spatial zones in one of which muscles are co-activated whereas in the adjacent zones, only one of the two antagonistic muscle groups is active (Fig. 3.25).

The notion of referent control helps solve both problems of action as well as some problems of perception or better to say, problems of action-perception coupling since it has been recognized for over a century that perception and action are interdependent (Helmholtz 1866). Previous attempts to solve action-perception problems that will be considered in this Chapter have been based on the efference copy (EC) concept introduced by von Holst and Mittelstaedt (von Holst and Mittelstaedt 1950; Holst 1954) or a similar concept of corollary discharges introduced by Sperry (1950). The feasibility of these concepts has been questioned, both on physical and physiological grounds (Chaps. 2 and 6). I will offer alternative solutions to some action-perception problems in the framework of referent control.

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.