Viscoelastic nanoscale properties of cuticle contribute to the high-pass properties of spider vibration receptor (Cupiennius saleiKeys)

Abstract
Atomic force microscopy (AFM) and surface force spectroscopy were applied in live spiders to their joint pad material located distal of the metatarsal lyriform organs, which are highly sensitive vibration sensors. The surface topography of the material is sufficiently smooth to probe the local nanomechanical properties with nanometre elastic deflections. Nanoscale loads were applied in the proximad direction on the distal joint region simulating the natural stimulus situation. The force curves obtained indicate the presence of a soft, liquid-like epicuticular layer (20–40 nm thick) above the pad material, which has much higher stiffness. The Young modulus of the pad material is close to 15 MPa at low frequencies, but increases rapidly with increasing frequencies approximately above 30 Hz to approximately 70 MPa at 112 Hz. The adhesive forces drop sharply by about 40% in the same frequency range. The strong frequency dependence of the elastic modulus indicates the viscoelastic nature of the pad material, its glass transition temperature being close to room temperature (25±2 °C) and, therefore, to its maximized energy absorption from low-frequency mechanical stimuli. These viscoelastic properties of the cuticular pad are suggested to be at least partly responsible for the high-pass characteristics of the vibration sensor's physiological properties demonstrated earlier.