Learning to stand with sensorimotor delays generalizes across directions and from hand to leg effectors.

Humans receive sensory information from the past, requiring the brain to overcome delays to perform daily motor skills such as standing upright. Because delays vary throughout the body and change over a lifetime, it would be advantageous to generalize learned control policies of balancing with delays across contexts. However, not all forms of learning generalize. Here, we use a robotic simulator to impose delays into human balance. When delays are imposed in one direction of standing, participants are initially unstable but relearn to balance by reducing the variability of their motor actions and transfer balance improvements to untrained directions. Upon returning to normal standing, aftereffects from learning are observed as small oscillations in control, yet they do not destabilize balance. Remarkably, when participants train to balance with delays using their hand, learning transfers to standing with the legs. Our findings establish that humans use experience to broadly update their neural control to balance with delays.

Vestibular control of deep and superficial lumbar muscles.

The active control of the lumbar musculature provides a stable platform critical for postures and goal-directed movements. Voluntary and perturbation-evoked motor commands can recruit individual lumbar muscles in a task-specific manner according to their presumed biomechanics. Here, we investigated the vestibular control of the deep and superficial lumbar musculature. Ten healthy participants were exposed to noisy electrical vestibular stimulation while balancing upright with their head facing forward, left, or right to characterize the differential modulation in the vestibular-evoked lumbar extensor responses in generating multidirectional whole body motion. We quantified the activation of the lumbar muscles on the right side using indwelling [deep multifidus, superficial multifidus, caudal longissimus (L4), and cranial longissimus (L1)] and high-density surface recordings. We characterized the vestibular-evoked responses using coherence and peak-to-peak cross-covariance amplitude between the vestibular and electromyographic signals. Participants exhibited responses in all lumbar muscles. The vestibular control of the lumbar musculature exhibited muscle-specific modulations: responses were larger in the longissimus (combined cranio-caudal) compared with the multifidus (combined deep-superficial) when participants faced forward (P < 0.001) and right (P = 0.011) but not when they faced left. The high-density surface recordings partly supported this observation: the location of the responses was more lateral when facing right compared with left (P < 0.001). The vestibular control of muscle subregions within the longissimus or the multifidus was similar. Our results demonstrate muscle-specific vestibular control of the lumbar muscles in response to perturbations of vestibular origin. The lack of differential activation of lumbar muscle subregions suggests the vestibular control of these subregions is co-regulated for standing balance.NEW & NOTEWORTHY We investigated the vestibular control of the deep and superficial lumbar extensor muscles using electrical vestibular stimuli. Vestibular stimuli elicited preferential activation of the longissimus muscle over the multifidus muscle. We did not observe clear regional activation of lumbar muscle subregions in response to the vestibular stimuli. Our findings show that the central nervous system can finely tune the vestibular control of individual lumbar muscles and suggest minimal regional variations in the activation of lumbar muscle subregions.

Real-world characterization of vestibular contributions during locomotion.

Introduction: The vestibular system, which encodes our head movement in space, plays an important role in maintaining our balance as we navigate the environment. While in-laboratory research demonstrates that the vestibular system exerts a context-dependent influence on the control of balance during locomotion, differences in whole-body and head kinematics between indoor treadmill and real-world locomotion challenge the generalizability of these findings. Thus, the goal of this study was to characterize vestibular-evoked balance responses in the real world using a fully portable system. Methods: While experiencing stochastic electrical vestibular stimulation (0-20 Hz, amplitude peak ± 4.5 mA, root mean square 1.25 mA) and wearing inertial measurement units (IMUs) on the head, low back, and ankles, 10 participants walked outside at 52 steps/minute (∼0.4 m/s) and 78 steps/minute (∼0.8 m/s). We calculated time-dependent coherence (a measure of correlation in the frequency domain) between the applied stimulus and the mediolateral back, right ankle, and left ankle linear accelerations to infer the vestibular control of balance during locomotion. Results: In all participants, we observed vestibular-evoked balance responses. These responses exhibited phasic modulation across the stride cycle, peaking during the middle of the single-leg stance in the back and during the stance phase for the ankles. Coherence decreased with increasing locomotor cadence and speed, as observed in both bootstrapped coherence differences (p < 0.01) and peak coherence (low back: 0.23 ± 0.07 vs. 0.16 ± 0.14, p = 0.021; right ankle: 0.38 ± 0.12 vs. 0.25 ± 0.10, p < 0.001; left ankle: 0.33 ± 0.09 vs. 0.21 ± 0.09, p < 0.001). Discussion: These results replicate previous in-laboratory studies, thus providing further insight into the vestibular control of balance during naturalistic movements and validating the use of this portable system as a method to characterize real-world vestibular responses. This study will help support future work that seeks to better understand how the vestibular system contributes to balance in variable real-world environments.