Statistically significant contrasts between EMG waveforms revealed using wavelet-based functional ANOVA.

We developed wavelet-based functional ANOVA (wfANOVA) as a novel approach for comparing neurophysiological signals that are functions of time. Temporal resolution is often sacrificed by analyzing such data in large time bins, increasing statistical power by reducing the number of comparisons. We performed ANOVA in the wavelet domain because differences between curves tend to be represented by a few temporally localized wavelets, which we transformed back to the time domain for visualization. We compared wfANOVA and ANOVA performed in the time domain (tANOVA) on both experimental electromyographic (EMG) signals from responses to perturbation during standing balance across changes in peak perturbation acceleration (3 levels) and velocity (4 levels) and on simulated data with known contrasts. In experimental EMG data, wfANOVA revealed the continuous shape and magnitude of significant differences over time without a priori selection of time bins. However, tANOVA revealed only the largest differences at discontinuous time points, resulting in features with later onsets and shorter durations than those identified using wfANOVA (P < 0.02). Furthermore, wfANOVA required significantly fewer (~1/4;×; P < 0.015) significant F tests than tANOVA, resulting in post hoc tests with increased power. In simulated EMG data, wfANOVA identified known contrast curves with a high level of precision (r(2) = 0.94 ± 0.08) and performed better than tANOVA across noise levels (P < <0.01). Therefore, wfANOVA may be useful for revealing differences in the shape and magnitude of neurophysiological signals (e.g., EMG, firing rates) across multiple conditions with both high temporal resolution and high statistical power.

Neuromechanical tuning of nonlinear postural control dynamics.

Postural control may be an ideal physiological motor task for elucidating general questions about the organization, diversity, flexibility, and variability of biological motor behaviors using nonlinear dynamical analysis techniques. Rather than presenting "problems" to the nervous system, the redundancy of biological systems and variability in their behaviors may actually be exploited to allow for the flexible achievement of multiple and concurrent task-level goals associated with movement. Such variability may reflect the constant "tuning" of neuromechanical elements and their interactions for movement control. The problem faced by researchers is that there is no one-to-one mapping between the task goal and the coordination of the underlying elements. We review recent and ongoing research in postural control with the goal of identifying common mechanisms underlying variability in postural control, coordination of multiple postural strategies, and transitions between them. We present a delayed-feedback model used to characterize the variability observed in muscle coordination patterns during postural responses to perturbation. We emphasize the significance of delays in physiological postural systems, requiring the modulation and coordination of both the instantaneous, "passive" response to perturbations as well as the delayed, "active" responses to perturbations. The challenge for future research lies in understanding the mechanisms and principles underlying neuromechanical tuning of and transitions between the diversity of postural behaviors. Here we describe some of our recent and ongoing studies aimed at understanding variability in postural control using physical robotic systems, human experiments, dimensional analysis, and computational models that could be enhanced from a nonlinear dynamics approach.

Mechanisms of motor adaptation in reactive balance control.

Balance control must be rapidly modified to provide stability in the face of environmental challenges. Although changes in reactive balance over repeated perturbations have been observed previously, only anticipatory postural adjustments preceding voluntary movements have been studied in the framework of motor adaptation and learning theory. Here, we hypothesized that adaptation occurs in task-level balance control during responses to perturbations due to central changes in the control of both anticipatory and reactive components of balance. Our adaptation paradigm consisted of a Training set of forward support-surface perturbations, a Reversal set of novel countermanding perturbations that reversed direction, and a Washout set identical to the Training set. Adaptation was characterized by a change in a motor variable from the beginning to the end of each set, the presence of aftereffects at the beginning of the Washout set when the novel perturbations were removed, and a return of the variable at the end of the Washout to a level comparable to the end of the Training set. Task-level balance performance was characterized by peak center of mass (CoM) excursion and velocity, which showed adaptive changes with repetitive trials. Only small changes in anticipatory postural control, characterized by body lean and background muscle activity were observed. Adaptation was found in the evoked long-latency muscular response, and also in the sensorimotor transformation mediating that response. Finally, in each set, temporal patterns of muscle activity converged towards an optimum predicted by a trade-off between maximizing motor performance and minimizing muscle activity. Our results suggest that adaptation in balance, as well as other motor tasks, is mediated by altering central sensitivity to perturbations and may be driven by energetic considerations.

A feedback model explains the differential scaling of human postural responses to perturbation acceleration and velocity.

Although the neural basis of balance control remains unknown, recent studies suggest that a feedback law on center-of-mass (CoM) kinematics determines the temporal patterning of muscle activity during human postural responses. We hypothesized that the same feedback law would also explain variations in muscle activity to support-surface translation as perturbation characteristics vary. Subject CoM motion was experimentally modulated using 34 different anterior-posterior support-surface translations of varying peak acceleration and velocity but the same total displacement. Electromyographic (EMG) recordings from several muscles of the lower limbs and trunk were compared to predicted EMG patterns from an inverted pendulum model under delayed feedback control. In both recorded and predicted EMG patterns, the initial burst of muscle activity scaled linearly with peak acceleration, whereas the tonic "plateau" region scaled with peak velocity. The relatively invariant duration of the initial burst was modeled by incorporating a transient, time-limited encoding of CoM acceleration inspired by muscle spindle primary afferent dynamic responses. The entire time course of recorded and predicted muscle activity compared favorably across all conditions, suggesting that the initial burst of muscle activity is not generated by feedforward neural mechanisms. Perturbation conditions were presented randomly and subjects maintained relatively constant feedback gains across all conditions. In contrast, an optimal feedback solution based on a trade-off between CoM stabilization and energy expenditure predicted that feedback gains should change with perturbation characteristics. These results suggest that an invariant feedback law was used to generate the entire time course of muscle activity across a variety of postural disturbances.

A feedback model reproduces muscle activity during human postural responses to support-surface translations.

Although feedback models have been used to simulate body motions in human postural control, it is not known whether muscle activation patterns generated by the nervous system during postural responses can also be explained by a feedback control process. We investigated whether a simple feedback law could explain temporal patterns of muscle activation in response to support-surface translations in human subjects. Previously, we used a single-link inverted-pendulum model with a delayed feedback controller to reproduce temporal patterns of muscle activity during postural responses in cats. We scaled this model to human dimensions and determined whether it could reproduce human muscle activity during forward and backward support-surface perturbations. Through optimization, we found three feedback gains (on pendulum acceleration, velocity, and displacement) and a common time delay that allowed the model to best match measured electromyographic (EMG) signals. For each muscle and each subject, the entire time courses of EMG signals during postural responses were well reconstructed in muscles throughout the lower body and resembled the solution derived from an optimal control model. In ankle muscles, >75% of the EMG variability was accounted for by model reconstructions. Surprisingly, >67% of the EMG variability was also accounted for in knee, hip, and pelvis muscles, even though motion at these joints was minimal. Although not explicitly required by our optimization, pendulum kinematics were well matched to subject center-of-mass (CoM) kinematics. Together, these results suggest that a common set of feedback signals related to task-level control of CoM motion is used in the temporal formation of muscle activity during postural control.