Cortical control during walking is most pronounced when the person is perturbed. Although seated locomotor tasks such as cycling or recumbent stepping improve walking performance, the electrocortical correlates of perturbed seated tasks have not been studied in detail. The primary purpose of this research was to quantify cortical and muscular responses to mechanical perturbations during recumbent stepping. We also aimed to quantify possible differences between young and older adults' responses to perturbed stepping. A secondary aim of this research was to determine the accuracy of electroencephalography (EEG) source imaging to interpret the electrocortical findings adequately. We hypothesized that both young and older adults would adapt to the perturbations by reducing their movement errors and reducing the anterior cingulate electrocortical activity. We also hypothesized that older adults would co-contract their agonist and antagonist muscles more than young adults in response to perturbations. Such stronger co-contraction would indicate older adults would have weaker corticomuscular connectivity in response to perturbations than young adults. Seventeen young adults and eleven older adults completed four perturbed arms and leg stepping tasks. We perturbed the stepping with brief 200ms increased movement resistance using a controllable servomotor on our recumbent stepper. We asked subjects to step smoothly, use both arms and legs and follow the visual pacing cue set at 60 steps per minute. We recorded brain activity with high-density EEG with 128 electrodes, muscular activity with 16 electromyography (EMG) sensors, and stepping kinematics using the servomotor's encoder. We quantified temporal and spatial motor errors from the stepping kinematics data. We used a novel post-processing approach to reject noise from EEG and estimated the electrocortical sources using independent component analysis and the current dipole estimation technique. We then performed a series of time-frequency analyses on the group EEG source data. We quantified EMG co-contraction for each of the perturbed and recovery steps. Finally, we used direct Directed Transfer Fucnction to determine the corticomuscular connectivity time-locked to young and older adults' perturbations. Quantifying the accuracy of source estimation showed that recording the three-dimensional EEG electrode locations would provide accurate source estimation up to a single Brodmann area. We also found that recording the precise location of the fiducials, i.e., the anatomical landmarks used to place the EEG electrodes, is critical for a reliable source estimation process. Motor errors did not show a reduction of errors with more perturbation experience for both young and older adults. Young adults showed significant theta-band (3-8 Hz) electrocortical activity locked to the perturbations at the anterior cingulate cortex, supplementary motor areas, and posterior parietal cortex. These locked spectral fluctuations decreased with more perturbation experience for the right-side perturbations and varied with perturbation timing. Older adults showed significant electrocortical activity with a wider spread of electrocortical sources in the motor and posterior parietal cortices. Older adults demonstrated more co-contracted muscle pairs than young adults, and co-contraction did not decrease with more perturbation experience. The results show that brief perturbation during recumbent stepping does not create error-based adaptation with reduced motor errors tied to more perturbation experience. However, these perturbations cause prolonged modifications in the motor patterns even after the perturbations are removed. Modulating the perturbation timing can tune both cortical activities at specific brain areas and modify muscular co-contraction behavior in older adults.


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Graduation Date





Huang, Helen


Doctor of Philosophy (Ph.D.)


College of Engineering and Computer Science


Mechanical and Aerospace Engineering

Degree Program

Mechanical Engineering




CFE0008531; DP0024207





Release Date

May 2021

Length of Campus-only Access


Access Status

Doctoral Dissertation (Open Access)