Indentation analysis of active viscoelastic microplasmodia of P. polycephalum

Simple organisms like Physarum polycephalum realize complex behavior, such as shortest path optimization or habituation, via mechanochemical processes rather than by a network of neurons. A full understanding of these phenomena requires detailed investigation of the underlying mechanical properties. To date, micromechanical measurements on P. polycephalum are sparse and lack reproducibility. This prompts study of microplasmodia, a reproducible and homogeneous form of P. polycephalum that resembles the plasmodial ectoplasm responsible for mechanical stability and generation of forces. We combine investigation of ultra-structure and dimension of P. polycephalum with the analysis of data obtained by indentation of microplasmodia, employing a novel nonlinear viscoelastic scaling model that accounts for finite dimension of the sample. We identify the multi-modal distribution of parameters such as Young's moduls, Poisson's ratio, and relaxation times associated with viscous processes that cover five orders of magnitude. Results suggest a characterization of microplasmodia as porous, compressible structures that act like elastic solids with high Young's modulus on short time scales, whereas on long time-scales and upon repeated indentation viscous behavior dominates and the effective modulus is significantly decreased. Furthermore, Young's modulus is found to oscillate in phase with shape of microplasmodia, emphasizing that modeling P. polycephalum oscillations as a driven oscillator with constant moduli is not practicable.