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Campaniform sensilla (CS) are mechanosensors embedded in the cuticle of insects. They are often found at locations near the joints of leg segments. On legs, CS are generally considered to respond directionally to cuticle bending during legged locomotion. It is currently unclear how CS locations affect strain levels at the CS, but this information is crucial for understanding how CS respond to stimuli. Here we present a parametric finite element model of the femoral CS field forDrosophilahind legs with 12 general and seven CS-specific parameters each. This model allows testing how changes in CS location, orientation and material property affect strain levels at each CS. We used experimentally acquired kinematic data and computed ground reaction forces to simulatein vivo-like forward stepping. The displacements found in this study at the physiological CS field location near the trochanter–femur joint are smaller than those necessary for conformation changes of ion channels involved in signal elicitation. Also, variation of material properties of the CS had little influence on displacement magnitudes at the CS cap where the sensory neuron attaches. Thus, our results indicate that ground reaction forces alone are unlikely to serve CS field activation during forward walking. 

Walking is the most common form of how animals move on land. The model organismDrosophila melanogasterhas become increasingly popular for studying how the nervous system controls behavior in general and walking in particular. Despite recent advances in tracking and modeling leg movements of walkingDrosophilain 3D, there are still gaps in knowledge about the biomechanics of leg joints due to the tiny size of fruit flies. For instance, the natural alignment of joint rotational axes was largely neglected in previous kinematic analyses. In this study, we therefore present a detailed kinematic leg model in which not only the segment lengths but also the main rotational axes of the joints were derived from anatomical landmarks, namely, the joint condyles. Our model with natural oblique joint axes is able to adapt to the 3D leg postures of straight and forward walking fruit flies with high accuracy. When we compared our model to an orthogonalized version, we observed that our model showed a smaller error as well as differences in the used range of motion (ROM), highlighting the advantages of modeling natural rotational axes alignment for the study of joint kinematics. We further found that the kinematic profiles of front, middle, and hind legs differed in the number of required degrees of freedom as well as their contributions to stepping, time courses of joint angles, and ROM. Our findings provide deeper insights into the joint kinematics of walking inDrosophila, and, additionally, will help to develop dynamical, musculoskeletal, and neuromechanical simulations. 

Insect load sensors, called campaniform sensilla (CS), measure strain changes within the cuticle of appendages. This mechanotransduction provides the neuromuscular system with feedback for posture and locomotion. Owing to their diverse morphology and arrangement, CS can encode different strain directions. We used nano-computed tomography and finite-element analysis to investigate how different CS morphologies within one location—the femoral CS field of the leg in the fruit fly Drosophila —interact under load. By investigating the influence of CS substructures' material properties during simulated limb displacement with naturalistic forces, we could show that CS substructures (i.e. socket and collar) influence strain distribution throughout the whole CS field. Altered socket and collar elastic moduli resulted in 5% relative differences in displacement, and the artificial removal of all sockets caused differences greater than 20% in cap displacement. Apparently, CS sockets support the distribution of distal strain to more proximal CS, while collars alter CS displacement more locally. Harder sockets can increase or decrease CS displacement depending on sensor location. Furthermore, high-resolution imaging revealed that sockets are interconnected in subcuticular rows. In summary, the sensitivity of individual CS is dependent on the configuration of other CS and their substructures.