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1. Investigating the role of low level reinforcement reflex loops in insect locomotion

   https://doi.org/10.1088/1748-3190/ac28ea  

   Goldsmith, C A ; Quinn, R D ; Szczecinski, N S ( October 2021 , Bioinspiration & Biomimetics)

   Abstract Insects are highly capable walkers, but many questions remain regarding how the insect nervous system controls locomotion. One particular question is how information is communicated between the ‘lower level’ ventral nerve cord (VNC) and the ‘higher level’ head ganglia to facilitate control. In this work, we seek to explore this question by investigating how systems traditionally described as ‘positive feedback’ may initiate and maintain stepping in the VNC with limited information exchanged between lower and higher level centers. We focus on the ‘reflex reversal’ of the stick insect femur-tibia joint between a resistance reflex (RR) and an active reaction in response to joint flexion, as well as the activation of populations of descending dorsal median unpaired (desDUM) neurons from limb strain as our primary reflex loops. We present the development of a neuromechanical model of the stick insect ( Carausius morosus ) femur-tibia (FTi) and coxa-trochanter joint control networks ‘in-the-loop’ with a physical robotic limb. The control network generates motor commands for the robotic limb, whose motion and forces generate sensory feedback for the network. We based our network architecture on the anatomy of the non-spiking interneuron joint control network that controls the FTi joint, extrapolated network connectivity based on known muscle responses,more »and previously developed mechanisms to produce ‘sideways stepping’. Previous studies hypothesized that RR is enacted by selective inhibition of sensory afferents from the femoral chordotonal organ, but no study has tested this hypothesis with a model of an intact limb. We found that inhibiting the network’s flexion position and velocity afferents generated a reflex reversal in the robot limb’s FTi joint. We also explored the intact network’s ability to sustain steady locomotion on our test limb. Our results suggested that the reflex reversal and limb strain reinforcement mechanisms are both necessary but individually insufficient to produce and maintain rhythmic stepping in the limb, which can be initiated or halted by brief, transient descending signals. Removing portions of this feedback loop or creating a large enough disruption can halt stepping independent of the higher-level centers. We conclude by discussing why the nervous system might control motor output in this manner, as well as how to apply these findings to generalized nervous system understanding and improved robotic control.« less
2. Computational Modeling of Spinal Locomotor Circuitry in the Age of Molecular Genetics

   https://doi.org/10.3390/ijms22136835  

   Ausborn, Jessica ; Shevtsova, Natalia A. ; Danner, Simon M. ( July 2021 , International Journal of Molecular Sciences)

   Neuronal circuits in the spinal cord are essential for the control of locomotion. They integrate supraspinal commands and afferent feedback signals to produce coordinated rhythmic muscle activations necessary for stable locomotion. For several decades, computational modeling has complemented experimental studies by providing a mechanistic rationale for experimental observations and by deriving experimentally testable predictions. This symbiotic relationship between experimental and computational approaches has resulted in numerous fundamental insights. With recent advances in molecular and genetic methods, it has become possible to manipulate specific constituent elements of the spinal circuitry and relate them to locomotor behavior. This has led to computational modeling studies investigating mechanisms at the level of genetically defined neuronal populations and their interactions. We review literature on the spinal locomotor circuitry from a computational perspective. By reviewing examples leading up to and in the age of molecular genetics, we demonstrate the importance of computational modeling and its interactions with experiments. Moving forward, neuromechanical models with neuronal circuitry modeled at the level of genetically defined neuronal populations will be required to further unravel the mechanisms by which neuronal interactions lead to locomotor behavior.
3. The Spring-Mass Model and Other Reductionist Models of Bipedal Locomotion on Inclines

   https://doi.org/10.1093/icb/icac047  

   Selvitella, Alessandro Maria ; Foster, Kathleen Lois ( June 2022 , Integrative and Comparative Biology)

   The spring-mass model is a model of locomotion aimed at giving the essential mathematical laws of the trajectory of the center of mass of an animal during bouncing gaits, such as hopping (one-dimensional) and running (two-dimensional). This reductionist mechanical system has been extensively investigated for locomotion over horizontal surfaces, whereas it has been largely neglected on other ecologically relevant surfaces, including inclines. For example, how the degree of inclination impacts the dynamics of the center of mass of the spring-mass model has not been investigated thoroughly. In this work, we derive a mathematical model which extends the spring-mass model to inclined surfaces. Among our results, we derive an approximate solution of the system, assuming a small angular sweep of the limb and a small spring compression during stance, and show that this approximation is very accurate, especially for small inclinations of the ground. Furthermore, we derive theoretical bounds on the difference between the Lagrangian and Lagrange equations of the true and approximate systems, and discuss locomotor stability questions of the approximate solutions. We test our models through a sensitivity analysis using parameters relevant to the locomotion of bipedal animals (quail, pheasant, guinea fowl, turkey, ostrich, and humans) and compare ourmore »approximate solution to the numerically derived solution of the exact system. We compare the two-dimensional spring-mass model on inclines with the one-dimensional spring-mass model to which it reduces under the limit of no horizontal velocity; we compare the two-dimensional spring-mass model on inclines with the inverted pendulum model on inclines towards which it converges in the case of high stiffness-to-mass ratio. We include comparisons with historically prevalent no-gravity approximations of these models, as well. The insights we have gleaned through all these comparisons and the ability of our approximation to replicate some of the kinematic changes observed in animals moving on different inclines (e.g., reduction in vertical oscillation of the center of mass and decreased stride length) underline the valuable and reasonable contributions that very simple, reductionist models, like the spring-mass model, can provide.

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4. Treadmill vs. overground walking: different response to physical interaction

   https://doi.org/10.1152/jn.00176.2017  

   Ochoa, Julieth ; Sternad, Dagmar ; Hogan, Neville ( October 2017 , Journal of Neurophysiology)

   Rehabilitation of human motor function is an issue of growing significance, and human-interactive robots offer promising potential to meet the need. For the lower extremity, however, robot-aided therapy has proven challenging. To inform effective approaches to robotic gait therapy, it is important to better understand unimpaired locomotor control: its sensitivity to different mechanical contexts and its response to perturbations. The present study evaluated the behavior of 14 healthy subjects who walked on a motorized treadmill and overground while wearing an exoskeletal ankle robot. Their response to a periodic series of ankle plantar flexion torque pulses, delivered at periods different from, but sufficiently close to, their preferred stride cadence, was assessed to determine whether gait entrainment occurred, how it differed across conditions, and if the adapted motor behavior persisted after perturbation. Certain aspects of locomotor control were exquisitely sensitive to walking context, while others were not. Gaits entrained more often and more rapidly during overground walking, yet, in all cases, entrained gaits synchronized the torque pulses with ankle push-off, where they provided assistance with propulsion. Furthermore, subjects entrained to perturbation periods that required an adaption toward slower cadence, even though the pulses acted to accelerate gait, indicating a neural adaptation ofmore »locomotor control. Lastly, during 15 post-perturbation strides, the entrained gait period was observed to persist more frequently during overground walking. This persistence was correlated with the number of strides walked at the entrained gait period (i.e., longer exposure), which also indicated a neural adaptation. NEW & NOTEWORTHY We show that the response of human locomotion to physical interaction differs between treadmill and overground walking. Subjects entrained to a periodic series of ankle plantar flexion torque pulses that shifted their gait cadence, synchronizing ankle push-off with the pulses (so that they assisted propulsion) even when gait cadence slowed. Entrainment was faster overground and, on removal of torque pulses, the entrained gait period persisted more prominently overground, indicating a neural adaptation of locomotor control.« less
5. Contribution of Afferent Feedback to Adaptive Hindlimb Walking in Cats: A Neuromusculoskeletal Modeling Study

   https://doi.org/10.3389/fbioe.2022.825149  

   Kim, Yongi ; Aoi, Shinya ; Fujiki, Soichiro ; Danner, Simon M. ; Markin, Sergey N. ; Ausborn, Jessica ; Rybak, Ilya A. ; Yanagihara, Dai ; Senda, Kei ; Tsuchiya, Kazuo ( April 2022 , Frontiers in Bioengineering and Biotechnology)

   Mammalian locomotion is generated by central pattern generators (CPGs) in the spinal cord, which produce alternating flexor and extensor activities controlling the locomotor movements of each limb. Afferent feedback signals from the limbs are integrated by the CPGs to provide adaptive control of locomotion. Responses of CPG-generated neural activity to afferent feedback stimulation have been previously studied during fictive locomotion in immobilized cats. Yet, locomotion in awake, behaving animals involves dynamic interactions between central neuronal circuits, afferent feedback, musculoskeletal system, and environment. To study these complex interactions, we developed a model simulating interactions between a half-center CPG and the musculoskeletal system of a cat hindlimb. Then, we analyzed the role of afferent feedback in the locomotor adaptation from a dynamic viewpoint using the methods of dynamical systems theory and nullcline analysis. Our model reproduced limb movements during regular cat walking as well as adaptive changes of these movements when the foot steps into a hole. The model generates important insights into the mechanism for adaptive locomotion resulting from dynamic interactions between the CPG-based neural circuits, the musculoskeletal system, and the environment.