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Free, publicly-accessible full text available December 13, 2025
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Abstract Studying the nervous system underlying animal motor control can shed light on how animals can adapt flexibly to a changing environment. We focus on the neural basis of feeding control in
Aplysia californica . Using the Synthetic Nervous System framework, we developed a model ofAplysia feeding neural circuitry that balances neurophysiological plausibility and computational complexity. The circuitry includes neurons, synapses, and feedback pathways identified in existing literature. We organized the neurons into three layers and five subnetworks according to their functional roles. Simulation results demonstrate that the circuitry model can capture the intrinsic dynamics at neuronal and network levels. When combined with a simplified peripheral biomechanical model, it is sufficient to mediate three animal-like feeding behaviors (biting, swallowing, and rejection). The kinematic, dynamic, and neural responses of the model also share similar features with animal data. These results emphasize the functional roles of sensory feedback during feeding.Free, publicly-accessible full text available May 20, 2025 -
The mechanical forces experienced during movement and the time constants of muscle activation are important determinants of the durations of behaviors, which may both be affected by size-dependent scaling. The mechanics of slow movements in small animals are dominated by elastic forces and are thus quasistatic (i.e., always near mechanical equilibrium). Muscular forces producing movement and elastic forces resisting movement should both scale identically (proportional to mass⅔), leaving the scaling of the time constant of muscle activation to play a critical role in determining behavioral duration. We tested this hypothesis by measuring the duration of feeding behaviors in the marine mollusc Aplysia californica whose body sizes spanned three orders of magnitude. The duration of muscle activation was determined by measuring the time it took for muscles to produce maximum force as Aplysia attempted to feed on tethered inedible seaweed, which provided an in vivo approximation of an isometric contraction. The timing of muscle activation scaled with mass0.3. The total duration of biting behaviors scaled identically, with mass0.3, indicating a lack of additional mechanical effects. The duration of swallowing behavior, however, exhibited a shallower scaling of mass0.17. We suggest that this was due to the allometric growth of the anterior retractor muscle during development, as measured by micro computed tomography scans (microCT) of buccal masses. Consequently, larger Aplysia did not need to activate their muscles as fully to produce equivalent forces. These results indicate that muscle activation may be an important determinant of the scaling of behavioral durations in quasistatic systems.
Free, publicly-accessible full text available April 8, 2025 -
This study introduces a novel neuromechanical model of rat hindlimbs with biarticular muscles producing walking movements without ground contact. The design of the control network is informed by the findings from our previous investigations into two-layer central pattern generators (CPGs). Specifically, we examined one plausible synthetic nervous system (SNS) designed to actuate 3 biarticular muscles, including the Biceps femoris posterior (BFP) and Rectus femoris (RF), both of which provide torque about the hip and knee joints. We conducted multiple perturbation tests on the simulation model to investigate the contribution of these two biarticular muscles in stabilizing perturbed hindlimb walking movements. We tested the BFP and RF muscles under three conditions: active, only passive tension, and fully disabled. Our results show that when these two biarticular muscles were active, they not only reduced the impact of external torques, but also facilitated rapid coordination of motion phases. As a result, the hindlimb model with biarticular muscles demonstrated faster recovery compared to our previous monoarticular muscle model.more » « less
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Abstract Soft earthworm‐like robots that exhibit mechanical compliance can, in principle, navigate through uneven terrains and constricted spaces that are inaccessible to traditional legged and wheeled robots. However, unlike the biological originals that they mimic, most of the worm‐like robots reported to date contain rigid components that limit their compliance, such as electromotors or pressure‐driven actuation systems. Here, a mechanically compliant worm‐like robot with a fully modular body that is based on soft polymers is reported. The robot is composed of strategically assembled, electrothermally activated polymer bilayer actuators, which are based on a semicrystalline polyurethane with an exceptionally large nonlinear thermal expansion coefficient. The segments are designed on the basis of a modified Timoshenko model, and finite element analysis simulation is used to describe their performance. Upon electrical activation of the segments with basic waveform patterns, the robot can move through repeatable peristaltic locomotion on exceptionally slippery or sticky surfaces and it can be oriented in any direction. The soft body enables the robot to wriggle through openings and tunnels that are much smaller than its cross‐section.
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Abstract Motor systems show an overall robustness, but because they are highly nonlinear, understanding how they achieve robustness is difficult. In many rhythmic systems, robustness against perturbations involves response of both the shape and the timing of the trajectory. This makes the study of robustness even more challenging. To understand how a motor system produces robust behaviors in a variable environment, we consider a neuromechanical model of motor patterns in the feeding apparatus of the marine mollusk Aplysia californica (Shaw et al. in J Comput Neurosci 38(1):25–51, 2015; Lyttle et al. in Biol Cybern 111(1):25–47, 2017). We established in (Wang et al. in SIAM J Appl Dyn Syst 20(2):701–744, 2021. https://doi.org/10.1137/20M1344974 ) the tools for studying combined shape and timing responses of limit cycle systems under sustained perturbations and here apply them to study robustness of the neuromechanical model against increased mechanical load during swallowing. Interestingly, we discover that nonlinear biomechanical properties confer resilience by immediately increasing resistance to applied loads. In contrast, the effect of changed sensory feedback signal is significantly delayed by the firing rates’ hard boundary properties. Our analysis suggests that sensory feedback contributes to robustness in swallowing primarily by shifting the timing of neural activation involved in the power stroke of the motor cycle (retraction). This effect enables the system to generate stronger retractor muscle forces to compensate for the increased load, and hence achieve strong robustness. The approaches that we are applying to understanding a neuromechanical model in Aplysia , and the results that we have obtained, are likely to provide insights into the function of other motor systems that encounter changing mechanical loads and hard boundaries, both due to mechanical and neuronal firing properties.more » « less
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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.more » « less
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Animal locomotion is influenced by a combination of constituent joint torques (e.g., due to limb inertia and passive viscoelasticity), which determine the necessary muscular response to move the limb. Across animal size-scales, the relative contributions of these constituent joint torques affect the muscular response in different ways. We used a multi-muscle biomechanical model to analyze how passive torque components change due to an animal’s size-scale during locomotion. By changing the size-scale of the model, we characterized emergent muscular responses at the hip as a result of the changing constituent torque profile. Specifically, we found that activation phases between extensor and flexor torques to be opposite between small and large sizes for the same kinematic motion. These results suggest general principles of how animal size affects neural control strategies. Our modeled torque profiles show a strong agreement with documented hindlimb torque during locomotion and can provide insights into the neural organization and muscle activation behavior of animals whose motion has not been extensively documented.more » « less