Axons span extreme distances and are subject to significant stretch deformations during limb movements or sudden head movements, especially during impacts. Yet, axon biomechanics, and its relation to the ultrastructure that allows axons to withstand mechanical stress, is poorly understood. Using a custom developed force apparatus, we demonstrate that chick dorsal root ganglion axons exhibit a tension buffering or strain-softening response, where its steady state elastic modulus decreases with increasing strain. We then explore the contributions from the various cytoskeletal components of the axon to show that the recently discovered membrane-associated actin-spectrin scaffold plays a prominent mechanical role. Finally, using a theoretical model, we argue that the actin-spectrin skeleton acts as an axonal tension buffer by reversibly unfolding repeat domains of the spectrin tetramers to release excess mechanical stress. Our results revise the current viewpoint that microtubules and their associated proteins are the only significant load-bearing elements in axons.

Spider silk possesses unique mechanical properties like large extensibility, high tensile strength, super-contractility, etc. Understanding these mechanical responses require characterization of the rheological properties of silk beyond the simple force-extension relations which are widely reported.Here we study the linear and non-linear viscoelastic properties of dragline silk obtained from social spiders Stegodyphus sarasinorum using a Micro-Extension Rheometer that we have developed.Unlike continuous extension data, our technique allows for the probing of the viscoelastic response by applying small perturbations about sequentially increasing steady-state strain values.In addition, we extend our analysis to obtain the characteristic stress relaxation times and the frequency responses of the viscous and elastic moduli. Using these methods, we show that in a small strain regime (0-4%) dragline silk of social spiders shows strain-softening response followed by strain-stiffening response at higher strains (> 4%). The stress relaxation time, on the other hand, increases monotonically with increasing strain for the entire range. We also show that silk stiffens while ageing within the typical lifetime of a web. Our results demand the inclusion of the kinetics of domain unfolding and refolding in the existing models to account for the relaxation time behaviour.

Rheological properties of a material often require to be probed under extensional deformation. Examples include fibrous materials such as spider-silk, high-molecular weight polymer melts, and the contractile response of living cells. Such materials have strong molecular-level anisotropies which are either inherent or are induced by an imposed extension. However, unlike shear rheology, which is well-established, techniques to perform extensional rheology are currently under development and setups are often custom-designed for the problem under study. In this article, we present a versatile device that can be used to conduct extensional deformation studies of samples at microscopic scales with simultaneous imaging. We discuss the operational features of this device and present a number of applications.

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