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Because soft animals are deformable, their locomotion is particularly affected by external forces and they are expected to face challenges controlling movements in different environments and orientations.We have used the caterpillar Manduca sexta to study neuromechanical strategies of soft-bodied scansorial locomotion. Manduca locomotion critically depends on the timing of proleg grip release, which is mediated by the principal planta retractor muscle and its single motoneuron, PPR. During upright crawling, PPR firing frequency increases approximately 0.6 s before grip release but during upsidedown crawling, this activity begins significantly earlier, possibly pretensioning the muscle. Under different loading conditions the timing of PPR activity changes relative to the stance/swing cycle. PPR motor activity is greater during upside-down crawling but these frequency changes are too small to produce significant differences in muscle force. Detailed observation of the proleg tip show that it swells before the retractor muscle is activated. This small movement is correlated with the activation of more posterior body segments, suggesting that it results from indirect mechanical effects. The timing and direction of this proleg displacement implies that proleg grip release is a dynamic interplay of mechanics and active neural control. [ABSTRACT FROM AUTHOR]

Aimed movements require that an animal accurately locates the target and correctly reaches that location. One such behavior is the defensive strike seen in Manduca sexta larva. These caterpillars respond to noxious mechanical stimuli applied to their abdomen with a strike of the mandibles towards the location of the stimulus. The accuracy with which the first strike movement reaches the stimulus site depends on the location of the stimulus. Reponses to dorsal stimuli are less accurate than those to ventral stimuli and the mandibles generally land ventral to the stimulus site. Responses to stimuli applied to anterior abdominal segments are less accurate than responses to stimuli applied to more posterior segments and the mandibles generally land posterior to the stimulus site. A trade-off between duration of the strike and radial accuracy is only seen in the anterior stimulus location (body segment A4). The lower accuracy of the responses to anterior and dorsal stimuli can be explained by the morphology of the animal; to reach these areas the caterpillar needs to move its body into a tight curve. Nevertheless, the accuracy is not exact in locations that the animal has shown it can reach, which suggests that consistently aiming more ventral and posterior of the stimulation site might be a defense strategy. [ABSTRACT FROM AUTHOR]

Abstract: The demand for measuring forces exerted by animals during locomotion has increased dramatically as biomechanists strive to understand and implement biomechanical control strategies. In particular, multi-axial force transducers are often required to capture animal limb coordination patterns. Most existing force transducers employ strain gages arranged in a Wheatstone bridge on a cantilever beam. Bi-axial measurements require duplicating this arrangement in the transverse direction. In this paper, we reveal a method to embed a Wheatstone bridge inside another to allow bi-axial measurements without additional strain gages or additional second beams. This hybrid configuration resolves two force components from a single bridge circuit and simplifies fabrication for the simultaneous assessment of normal and transverse loads. This design can be implemented with two-dimensional fabrication techniques and can even be used to modify a common full bridge cantilever force transducer. As a demonstration of the new design, we built a simple beam which achieved bi-axial sensing capability that outperformed a conventional half-bridge-per-axis bi-axial strain gage design. We have used this design to measure the ground reaction forces of a crawling caterpillar and a caterpillar-mimicking soft robot. The simplicity and increased sensitivity of this method could facilitate bi-axial force measurements for experimental biologists. [Copyright &y& Elsevier]

Caterpillars can increase their body mass 10,000-fold in 2weeks. It is therefore remarkable that most caterpillars appear to maintain the same locomotion kinematics throughout their entire larval stage. This study examined how the body properties of a caterpillar might change to accommodate such dramatic changes in body load. Using Manduca sexta as a model system, we measured changes in body volume, tissue density and baseline body pressure, and the dimensions of load-bearing tissues (the cuticle and muscles) over a body mass range from milligrams to several grams. All Manduca biometrics relevant to the hydrostatic skeleton scaled allometrically but close to the isometric predictions. Body density and pressure were almost constant. We next investigated the effects of scaling on the bending stiffness of the caterpillar hydrostatic skeleton. The anisotropic non- linear mechanical response of Manduca muscles and soft cuticle has previously been quantified and modeled with constitutive equations. Using biometric data and these material laws, we constructed finite element models to simulate a hydrostatic skeleton under different conditions. The results show that increasing the internal pressure leads to a non-linear increase in bending stiffness. Increasing the body size results in a decrease in the normalized bending stiffness. Muscle activation can double this stiffness in the physiological pressure range, but thickening the cuticle or increasing the muscle area reduces the structural stiffness. These non-linear effects may dictate the effectiveness of a hydrostatic skeleton at different sizes. Given the shared anatomy and size variation in Lepidoptera larvae, these mechanical scaling constraints may implicate the diverse locomotion strategies in different species. [ABSTRACT FROM AUTHOR]

Abstract: The mechanical properties of soft tissues are important for the control of motion in many invertebrates. Pressurized cylindrical animals such as worms have circumferential reinforcement of the body wall; however, no experimental characterization of comparable anisotropy has been reported for climbing larvae such as caterpillars. Using uniaxial, real-time fluorescence extensometry on millimeter scale cuticle specimens we have quantified differences in the mechanical properties of cuticle to circumferentially and longitudinally applied forces. Based on these results and the composite matrix–fiber structure of cuticle, a pseudo-elastic transversely isotropic constitutive material model was constructed with circumferential reinforcement realized as a Horgan–Saccomandi strain energy function. This model was then used numerically to describe the anisotropic material properties of Manduca cuticle. The constitutive material model will be used in a detailed finite-element analysis to improve our understanding of the mechanics of caterpillar crawling. [Copyright &y& Elsevier]