Elasticity in Ionically Cross-Linked Neurofilament Networks

Neurofilaments are found in abundance in the cytoskeleton of neurons, where they act as an intracellular framework protecting the neuron from external stresses. To elucidate the nature of the mechanical properties that provide this protection, we measure the linear and nonlinear viscoelastic properties of networks of neurofilaments. These networks are soft solids that exhibit dramatic strain stiffening above critical strains of 30-70{\%}. Surprisingly, divalent ions, such as Mg$^{2+}$, Ca$^{2+}$, and Zn$^{2+}$, act as effective cross-linkers for neurofilament networks, controlling their solid-like elastic response. This behavior is comparable to that of actin-binding proteins in reconstituted filamentous actin. We show that the elasticity of neurofilament networks is entropic in origin and is consistent with a model for cross-linked semiflexible networks, which we use to quantify the cross-linking by divalent ions. Ultimately, we are able to extract microstructural network parameters such as the persistence length and the average distance between cross-links directly from bulk rheology.