Experts, but not novices, exhibit StartReact indicating experts use the reticulospinal system more than novices.

Motor skill acquisition utilizes a wide array of neural structures; however, few articles evaluate how the relative contributions of these structures shift over the course of learning. Recent evidence from rodents and songbirds suggests there is a transfer from cortical to subcortical structures following intense, repetitive training. Evidence from humans indicate that the reticulospinal system is modulated over the course of skill acquisition and may be a subcortical facilitator of learning. The objective of this study was to evaluate how reticulospinal contributions are modulated by task expertise. Reticulospinal contributions were assessed using StartReact (SR). We hypothesized that expert typists would show SR during an individuated, keystroke task but SR would be absent in novices. Expert (75.2 ± 9.8 WPM) and novice typists (41.6 ± 8.2 WPM) were evaluated during an individuated, keystroke movements. In experts, SR was present but was absent in novices. Together, these results suggest that experts use reticulospinal contributions more for movement than novices indicating that the reticular formation becomes increasingly important for movement execution in highly trained, skilled tasks even those that require individuated movement of the fingers.

Startle evokes nearly identical movements in multi-jointed, two-dimensional reaching tasks.

StartReact is the ability of the startle reflex to involuntarily release a planned movement in the presence of a loud acoustic stimulus resulting in muscle activity patterns and kinematics that are tightly regulated and scaled with the intended action. Previous studies demonstrated startReact's robustness during simple single-joint reaching tasks and found no difference between startReact and voluntary movements for movement kinematics and muscle activation patterns. However, startReact has not been evaluated during multi-joint reaching movements with multiple degrees of freedom. It is unclear if startReact would evoke accurate and precise multi-joint reaching movements in an unrestricted workspace. Furthermore, if tested more rigorously, multi-joint startReact movement kinematics and muscle activation patterns might not be truly equivalent despite showing no difference through traditional ANOVAs. A previous study found multi-joint startReact was possible during unrestricted elbow and shoulder movement when reaching to a forward target. Therefore, we hypothesized that startReact would evoke similar multi-joint reaching movements for movement accuracy and muscle activation patterns when compared to voluntary movements in a multi-directional workspace. Expanding upon the previous study, our study uses a larger workspace and fully evaluates movement kinematics and muscle activations patterns. Results confirmed our hypothesis and found startReact movements were readily evoked in all directions. StartReact responses presented stereotypically earlier muscle activation, but the relative timing of agonist/antagonist firing pairs between startReact and voluntary movements remained similar. Results demonstrate that startReact is robustly present and equivalent in multi-joint reaching tasks and has potential clinical use for evaluating healthy and impaired movement.

Evidence for startle as a measurable behavioral indicator of motor learning.

The ability of the classic startle reflex to evoke voluntarily prepared movement involuntarily has captured the attention of neuroscientists for its wide-ranging functional utility and potential uses in patient populations. To date, there is only one documented task resistant to the startReact phenomenon-index finger abduction. Previous reports have suggested the lack of startReact is due to different neural mechanisms driving individuated finger movement and more proximal joint control (e.g. elbow, wrist movement). However, an alternative hypothesis exists. Though not particularly difficult to execute, isolated index finger abduction is rarely performed during activities of daily living and is not a natural correlate to common individuated finger tasks. We propose that startReact can be evoked during individuated finger movements but only during tasks that are highly trained or familiar. The objective of this study was to determine the impact of a 2-week training regimen on the ability to elicit startReact. We found evidence in support of our hypothesis that following training, individuated movements of the hands (specifically index finger abduction) become susceptible to startReact. This is significant not only because it indicates that individuated finger movements are in fact amenable to startReact, but also that startle has differential response characteristics in novel tasks compared to highly trained tasks suggesting that startle is a measurable behavioral indicator of motor learning.

Instruction-dependent modulation of the long-latency stretch reflex is associated with indicators of startle.

Long-latency responses elicited by postural perturbation are modulated by how a subject is instructed to respond to the perturbation, yet the neural pathways responsible for this modulation remain unclear. The goal of this study was to determine whether instruction-dependent modulation is associated with activity in brainstem pathways contributing to startle. Our hypothesis was that elbow perturbations can evoked startle, indicated by activity in the sternocleidomastoid muscle (SCM). Perturbation responses were compared to those elicited by a loud acoustic stimulus, known to elicit startle. Postural perturbations and startling acoustic stimuli both evoked SCM activity, but only when a ballistic elbow extension movement was planned. Both stimuli triggered SCM activity with the same probability. When SCM activity was present, there was an associated early onset of triceps electromyographic (EMG), as required for the planned movement. This early EMG onset occurred at a time often attributed to long-latency stretch reflexes (75-100 ms). The nature of the perturbation-triggered EMG (excitatory or inhibitory) was independent of the perturbation direction (flexion or extension) indicating that it was not a feedback response appropriate for returning the limb to its original position. The net EMG response to perturbations delivered after a movement had been planned could be explained as the sum of a stretch reflex opposing the perturbation and a startle-evoked response associated with the prepared movement. These results demonstrate that rapid perturbations can trigger early release of a planned ballistic movement, and that this release is associated with activity in the brainstem pathways contributing to startle reflexes.

Contributions of altered stretch reflex coordination to arm impairments following stroke.

Patterns of stereotyped muscle coactivation, clinically referred to as synergies, emerge following stroke and impair arm function. Although researchers have focused on cortical contributions, there is growing evidence that altered stretch reflex pathways may also contribute to impairment. However, most previous reflex studies have focused on passive, single-joint movements without regard to their coordination during volitional actions. The purpose of this study was to examine the effects of stroke on coordinated activity of stretch reflexes elicited in multiple arm muscles following multijoint perturbations. We hypothesized that cortical injury results in increased stretch reflexes of muscles characteristic of the abnormal flexor synergy during active arm conditions. To test this hypothesis, we used a robot to apply position perturbations to impaired arms of 10 stroke survivors and dominant arms of 8 healthy age-matched controls. Corresponding reflexes were assessed during volitional contractions simulating different levels of gravitational support, as well as during voluntary flexion and extension of the elbow and shoulder. Reflexes were quantified by average rectified surface electromyogram, recorded from eight muscles spanning the elbow and shoulder. Reflex coordination was quantified using an independent components analysis. We found stretch reflexes elicited in the stroke group were significantly less sensitive to changes in background muscle activation compared with those in the control group (P < 0.05). We also observed significantly increased reflex coupling between elbow flexor and shoulder abductor-extensor muscles in stroke subjects relative to that in control subjects. This increased coupling was present only during volitional tasks that required elbow flexion (P < 0.001), shoulder extension (P < 0.01), and gravity opposition (P < 0.01), but not during the "no load" condition. During volitional contractions, reflex amplitudes scaled with the level of impairment, as assessed by Fugl-Meyer scores (r(2) = 0.63; P < 0.05). We conclude that altered reflex coordination is indicative of motor impairment level and may contribute to impaired arm function following stroke.

Interactions between limb and environmental mechanics influence stretch reflex sensitivity in the human arm.

Stretch reflexes contribute to arm impedance and longer-latency stretch reflexes exhibit increased sensitivity during interactions with compliant or unstable environments. This increased sensitivity is consistent with a regulation of arm impedance to compensate for decreased stability of the environment, but the specificity of this modulation has yet to be investigated. Many tasks, such as tool use, compromise arm stability along specific directions, and stretch reflexes tuned to those directions could present an efficient mechanism for regulating arm impedance in a task-appropriate manner. To be effective, such tuning should adapt not only to the mechanical properties of the environment but to those properties in relation to the arm, which also has directionally specific mechanical properties. The purpose of this study was to investigate the specificity of stretch reflex modulation during interactions with mechanical environments that challenge arm stability. The tested environments were unstable, having the characteristics of a negative stiffness spring. These were either aligned or orthogonal to the direction of maximal endpoint stiffness for each subject. Our results demonstrate preferential increases in reflexes, elicited within 50-100 ms of perturbation onset, to perturbations applied specifically along the direction of the destabilizing environments. This increase occurred only when the magnitude of the environmental instability exceeded endpoint stiffness along the same direction. These results are consistent with task-specific reflex modulation tuned to the mechanical properties of the environment relative to those of the human arm. They demonstrate a highly adaptable, involuntary mechanism that may be used to modulate limb impedance along specific directions.

Mechanical perturbations applied during impending movement evoke startle-like responses.

Stretch reflexes have been considered one of the simplest circuits in the human nervous system. Yet, their role is controversial given that they assist or resist an imposed perturbation depending on the task instruction. Evidence shows that a loud acoustic stimulus applied prior to an impending movement elicits a movement-direction dependent muscle activity. In our study, we found that a perturbation can also trigger this early onset of movement, if applied during movement preparation. These responses were also perturbation direction dependent. This suggests an interaction of between the limb-stabilizing stretch reflexes and the voluntary activity.

Reflex modulation is linked to the orientation of arm mechanics relative to the environment.

To successfully complete a motor task, it is necessary to control not only the kinematics and dynamics of a limb, but also its mechanical properties. In a multijoint task such as the control of arm posture, limb mechanics are directional, resisting external disturbances more effectively in certain directions than others. It has been demonstrated that feedforward neuromotor pathways can regulate these directional characteristics of the arm to compensate for changes in the mechanical properties of the environment. However, it is unclear if spinal reflex pathways exhibit a similar specificity. The present results suggest that the sensitivity of the human stretch reflex also can be tuned to adapt the mechanical properties of the arm in a task appropriate manner. We hypothesized that the orientation of arm mechanics relative to the mechanical properties of the environment would influence reflex adaptation. Two destabilizing environments, oriented relative to the mechanical properties of the arm, were used to test this hypothesis. These environments were simulated using a 3 degrees of freedom (DOF) robot, which also was used to perturb arm posture. The resulting reflexes, assessed by electromyograms recorded from 8 muscles, were found to modulate in accordance with how the environmental instability was oriented relative to the mechanical properties of the arm. Our results suggest that stretch sensitive reflexes throughout the arm are modulated in a coordinated manner corresponding to the orientation of arm mechanics relative to the environment.

Altered multijoint reflex coordination is indicative of motor impairment level following stroke.

Following stroke, individuals often are unable to activate their elbow and shoulder muscles independently. There is growing evidence that altered reflex pathways may contribute to these abnormal patterns of activation or muscle synergies. Most studies investigating reflex function following stroke have examined only individual joints at rest. Thus, the purpose of this study was to quantify multijoint reflex contributions to the stereotyped muscle synergies commonly observed following stroke. We hypothesized that the patterns of reflex coordination mirror the abnormal muscle coactivity patterns previously reported for voluntary activation. 10 chronic stroke and 8 age-matched control subjects participated. Reflexes were elicited by perturbing the arm with a 3 degree of freedom robot while subjects exerted voluntary forces at the elbow and shoulder. The force conditions tested were selected to assess the influence of gravity and the influence of joint torque generation without gravity on reflex coordination. Reflex magnitude was quantified by the average rectified electromyogram, recorded from 8 muscles that span the elbow and shoulder. Patterns of reflex coordination were quantified using independent components analysis. Results show significant reflex coupling between elbow flexor and shoulder abductor-extensor muscles in stroke patients during isolated elbow and shoulder torque generation and during active arm support against gravity. Identified patterns of stretch reflex coordination were consistent with the stereotyped voluntary flexion synergy, suggesting reflex pathways contribute to abnormal muscle coordination following stroke.

Nonparametric identification of the elbow joint stiffness under compliant loads.

Several nonparametric system identification techniques have been used to estimate the dynamic joint stiffness of the human elbow. Most studies involved a very stiff environment, but some studies have also shown that stiffness is modified in response to environmental compliance. However, using the same identification technique used under very stiff conditions to do identification under compliant conditions leads to a biased estimate. This is due to the presence of feedback in the latter. In this paper, we use a nonparametric identification algorithm to demonstrate this problem. We then show how instrumental variables can be employed to obtain an unbiased estimate of the same. Both simulations as well as experimental data are used to this effect.