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.
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Biomechanical and Sensory Feedback Regularize the Behavior of Different Locomotor Central Pattern Generators
This work presents an in-depth numerical investigation into a hypothesized two-layer central pattern generator (CPG) that controls mammalian walking and how different parameter choices might affect the stepping of a simulated neuromechanical model. Particular attention is paid to the functional role of features that have not received a great deal of attention in previous work: the weak cross-excitatory connectivity within the rhythm generator and the synapse strength between the two layers. Sensitivity evaluations of deafferented CPG models and the combined neuromechanical model are performed. Locomotion frequency is increased in two different ways for both models to investigate whether the model’s stability can be predicted by trends in the CPG’s phase response curves (PRCs). Our results show that the weak cross-excitatory connection can make the CPG more sensitive to perturbations and that increasing the synaptic strength between the two layers results in a trade-off between forced phase locking and the amount of phase delay that can exist between the two layers. Additionally, although the models exhibit these differences in behavior when disconnected from the biomechanical model, these differences seem to disappear with the full neuromechanical model and result in similar behavior despite a variety of parameter combinations. This indicates that the neural variables do not have to be fixed precisely for stable walking; the biomechanical entrainment and sensory feedback may cancel out the strengths of excitatory connectivity in the neural circuit and play a critical role in shaping locomotor behavior. Our results support the importance of including biomechanical models in the development of computational neuroscience models that control mammalian locomotion.
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- Award ID(s):
- 2015317
- PAR ID:
- 10423851
- Date Published:
- Journal Name:
- Biomimetics
- Volume:
- 7
- Issue:
- 4
- ISSN:
- 2313-7673
- Page Range / eLocation ID:
- 226
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Synopsis The central pattern generator (CPG) in anguilliform swimming has served as a model for examining the neural basis of locomotion. This system has been particularly valuable for the development of mathematical models. As our biological understanding of the neural basis of locomotion has expanded, so too have these models. Recently, there have been significant advancements in our understanding of the critical role that mechanosensory feedback plays in robust locomotion. This work has led to a push in the field of mathematical modeling to incorporate mechanosensory feedback into CPG models. In this perspective piece, we review advances in the development of these models and discuss how newer complex models can support biological investigation. We highlight lamprey spinal cord regeneration as an area that can both inform these models and benefit from them.
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ABSTRACT Balitorid loaches are a family of fishes that exhibit morphological adaptations to living in fast flowing water, including an enlarged sacral rib that creates a ‘hip’-like skeletal connection between the pelvis and the axial skeleton. The presence of this sacral rib, the robustness of which varies across the family, is hypothesized to facilitate terrestrial locomotion seen in the family. Terrestrial locomotion in balitorids is unlike that of any known fish: the locomotion resembles that of terrestrial tetrapods. Emergence and convergence of terrestrial locomotion from water to land has been studied in fossils; however, studying balitorid walking provides a present-day natural laboratory to examine the convergent evolution of walking movements. We tested the hypothesis that balitorid species with more robust connections between the pelvic and axial skeleton (M3 morphotype) are more effective at walking than species with reduced connectivity (M1 morphotype). We predicted that robust connections would facilitate travel per step and increase mass support during movement. We collected high-speed video of walking in seven balitorid species to analyze kinematic variables. The connection between internal anatomy and locomotion on land are revealed herein with digitized video analysis, μCT scans, and in the context of the phylogenetic history of this family of fishes. Our species sampling covered the extremes of previously identified sacral rib morphotypes, M1 and M3. Although we hypothesized the robustness of the sacral rib to have a strong influence on walking performance, there was not a large reduction in walking ability in the species with the least modified rib (M1). Instead, walking kinematics varied between the two balitorid subfamilies with a generally more ‘walk-like’ behavior in the Balitorinae and more ‘swim-like’ behavior in the Homalopteroidinae. The type of terrestrial locomotion displayed in balitorids is unique among living fishes and aids in our understanding of the extent to which a sacral connection facilitates terrestrial walking.more » « less
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- Free Publicly Accessible Full Text
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.3390/biomimetics7040226
