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Abstract To understand how behaviors arise in animals, it is necessary to investigate both the neural circuits and the biomechanics of the periphery. A tractable model system for studying multifunctional control is the feeding apparatus of the marine molluskAplysia californica. Previousin silicoandin robotomodels have investigated how the nervous and muscular systems interact in this system. However, these models are still limited in their ability to matchin vivodata both qualitatively and quantitatively. We introduce a new neuromechanical model ofAplysiafeeding that combines a modified version of a previously developed neural model with a novel biomechanical model that better reflects the anatomy and kinematics ofAplysiafeeding. The model was calibrated using a combination of previously measured biomechanical parameters and hand-tuning to behavioral data. Using this model, simulated feeding experiments were conducted, and the resulting behavioral metrics were compared to animal data. The model successfully produces three key behaviors seen inAplysiaand demonstrates a good quantitative agreement with biting and swallowing behaviors. Additional work is needed to match rejection behavior quantitatively and to reflect qualitative observations related to the relative contributions of two key muscles, the hinge and I3. Future improvements will focus on incorporating the effects of deformable 3D structures in the simulated buccal mass. 

Resilin, an elastomeric protein with remarkable physical properties that outperforms synthetic rubbers, is a near-ubiquitous feature of the power amplification mechanisms used by jumping insects. Catapult-like mechanisms, which incorporate elastic energy stores formed from a composite of stiff cuticle and resilin, are frequently used by insects to translate slow muscle contractions into rapid-release recoil movements. The precise role of resilin in these jumping mechanisms remains unclear, however. We used RNAi to reduce resilin deposition in the principal energy-storing springs of the desert locust (Schistocerca gregaria) before measuring jumping performance. Knockdown reduced the amount of resilin-associated fluorescence in the semilunar processes (SLPs) by 44% and reduced the cross-sectional area of the tendons of the hind leg extensor-tibiae muscle by 31%. This affected jumping in three ways: First, take-off velocity was reduced by 15% in knockdown animals, which could be explained by a change in the extrinsic stiffness of the extensor-tibiae tendon caused by the decrease in its cross-sectional area. Second, knockdown resulted in permanent breakages in the hind legs of 29% of knockdown locusts as tested by electrical stimulation of the extensor muscle, but none in controls. Third, knockdown locusts exhibited a greater decline in distance jumped when made to jump in rapid succession than did controls. We conclude that stiff cuticle acts as the principal elastic energy store for insect jumping, while resilin protects these more brittle structures against breakage from repeated use. 

Abstract There is increasing interest from evolutionary biologists in the evolution of avian bill shape, how the bill is used during feeding and, in particular, the bite forces the bill can deliver. Bite force exhibits isometry with the total mass of the jaw musculature, but there is variation in the functional categories of the jaw muscles in different avian taxa. Qualitative descriptions of the jaw musculature do not allow analysis of the relative contributions that adductor or retractor muscles play in generating a bite force. This study is a meta-analysis of published data for body mass and the mass of the jaw musculature in 66 bird species from 10 orders. The masses of the different muscles contributing to adduction and retraction in closing the jaw, and to depression and protraction in opening the jaw, were summed and allometric relationships explored before investigating the effects of taxonomic order on these relationships. The categories of muscles, and the masses of each category of jaw musculature varied among avian orders. Some species, such as the flightless ratites, had relatively small jaw muscle mass but parrots had an additional adductor muscle. Phylogenetically controlled relationships between body mass and the mass of each muscle category irrespective of taxonomic order were isometric. However, analysis of covariance revealed significant interactions between body mass and taxonomic order. Most orders had low values for body-mass-specific muscle masses in the jaw with the notable exceptions of the Passeriformes (songbirds) and Psittaciformes (parrots). The values of these orders were 3–4 times greater, although the relative amounts of muscles contributing to adduction and retraction were similar in Psittaciformes, but adduction was markedly higher in Passeriformes. The results of these analyses highlight the lack of species-specific data for most birds, which is adversely impacting our understanding of the anatomical features that are determining the functional properties of the bill during feeding. 

Abstract The coordination of complex behavior requires knowledge of both neural dynamics and the mechanics of the periphery. The feeding system ofAplysia californicais an excellent model for investigating questions in soft body systems’ neuromechanics because of its experimental tractability. Prior work has attempted to elucidate the mechanical properties of the periphery by using a Hill-type muscle model to characterize the force generation capabilities of the key protractor muscle responsible for movingAplysia’s grasper anteriorly, the I2 muscle. However, the I1/I3 muscle, which is the main driver of retractions ofAplysia’s grasper, has not been characterized. Because of the importance of the musculature’s properties in generating functional behavior, understanding the properties of muscles like the I1/I3 complex may help to create more realistic simulations of the feeding behavior ofAplysia, which can aid in greater understanding of the neuromechanics of soft-bodied systems. To bridge this gap, in this work, the I1/I3 muscle complex was characterized using force-frequency, length-tension, and force-velocity experiments and showed that a Hill-type model can accurately predict its force-generation properties. Furthermore, the muscle’s peak isometric force and stiffness were found to exceed those of the I2 muscle, and these results were analyzed in the context of prior studies on the I1/I3 complex’s kinematics in vivo. 

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 inAplysia californica. Using the Synthetic Nervous System framework, we developed a model ofAplysiafeeding 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. 

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. 

Abstract The Orthoptera are a diverse insect order well known for their locomotive capabilities. To jump, the bush-cricket uses a muscle actuated (MA) system in which leg extension is actuated by contraction of the femoral muscles of the hind legs. In comparison, the locust uses a latch mediated spring actuated (LaMSA) system, in which leg extension is actuated by the recoil of spring-like structure in the femur. The aim of this study was to describe the jumping kinematics ofMecopoda elongata(Tettigoniidae) and compare this to existing data inSchistocerca gregaria(Acrididae), to determine differences in control of rotation during take-off between similarly sized MA and LaMSA jumpers. 269 jumps from 67 individuals ofM. elongatawith masses from 0.014 g to 3.01 g were recorded with a high-speed camera setup. InM. elongata, linear velocity increased with mass^0.18and the angular velocity (pitch) decreased with mass^−0.13. InS. gregaria, linear velocity is constant and angular velocity decreases with mass^−0.24. Despite these differences in velocity scaling, the ratio of translational kinetic energy to rotational kinetic energy was similar for both species. On average, the energy distribution ofM. elongatawas distributed 98.8% to translational kinetic energy and 1.2% to rotational kinetic energy, whilst inS. gregariait is 98.7% and 1.3%, respectively. This energy distribution was independent of size for both species. Despite having two different jump actuation mechanisms, the ratio of translational and rotational kinetic energy formed during take-off is fixed across these distantly related orthopterans. 

Abstract Locusts ( Schistocerca gregaria ) jump using a latch mediated spring actuated system in the femur-tibia joint of their metathoracic legs. These jumps are exceptionally fast and display angular rotation immediately after take-off. In this study, we focus on the angular velocity, at take-off, of locusts ranging between 0.049 and 1.50 g to determine if and how rotation-rate scales with size. From 263 jumps recorded from 44 individuals, we found that angular velocity scales with mass −0.33 , consistent with a hypothesis of locusts having a constant rotational kinetic energy density. Within the data from each locust, angular velocity increased proportionally with linear velocity, suggesting the two cannot be independently controlled and thus a fixed energy budget is formed at take-off. On average, the energy budget of a jump is distributed 98.7% to translational kinetic energy and gravitational potential energy, and 1.3% to rotational kinetic energy. The percentage of energy devoted to rotation was constant across all sizes of locusts and represents a very small proportion of the energy budget. This analysis suggests that smaller locusts find it harder to jump without body rotation. 

ABSTRACT Small organisms use propulsive springs rather than muscles to repeatedly actuate high acceleration movements, even when constrained to tiny displacements and limited by inertial forces. Through integration of a large kinematic dataset, measurements of elastic recoil, energetic math modeling and dynamic math modeling, we tested how trap-jaw ants (Odontomachus brunneus) utilize multiple elastic structures to develop ultrafast and precise mandible rotations at small scales. We found that O. brunneus develops torque on each mandible using an intriguing configuration of two springs: their elastic head capsule recoils to push and the recoiling muscle–apodeme unit tugs on each mandible. Mandibles achieved precise, planar, circular trajectories up to 49,100 rad s−1 (470,000 rpm) when powered by spring propulsion. Once spring propulsion ended, the mandibles moved with unconstrained and oscillatory rotation. We term this mechanism a ‘dual spring force couple’, meaning that two springs deliver energy at two locations to develop torque. Dynamic modeling revealed that dual spring force couples reduce the need for joint constraints and thereby reduce dissipative joint losses, which is essential to the repeated use of ultrafast, small systems. Dual spring force couples enable multifunctionality: trap-jaw ants use the same mechanical system to produce ultrafast, planar strikes driven by propulsive springs and for generating slow, multi-degrees of freedom mandible manipulations using muscles, rather than springs, to directly actuate the movement. Dual spring force couples are found in other systems and are likely widespread in biology. These principles can be incorporated into microrobotics to improve multifunctionality, precision and longevity of ultrafast systems.