Symmetry breaking transition towards directional locomotion in Physarum microplasmodia

True slime mold Physarum polycephalum has been widely used as a model organism to study flow-driven amoeboid locomotion as well as the dynamics of its complex mechanochemical self-oscillations. The aim of this work is to quantify the mechanical aspects of symmetry breaking and its transition into directional flow-driven amoeboid locomotion in small (<∼ 200 µm) fragments of Physarum polycephalum. To this end, we combined measurements of traction stresses, fragment morphology, and ectoplasmic microrheology with experimental manipulations of cell-substrate adhesion, cortical strength and microplasmodium size. These measurements show that initiation of locomotion is accompanied by the symmetry breaking of traction stresses and the polarization of ectoplasmic mechanical properties, with the rear part of the microplasmodium becoming significantly stiffer after the onset of locomotion. Our experimental data suggests that the initiation of locomotion in Physarum could be analogous to an interfacial instability process and that microplasmodial size is a critical parameter governing the instability. Specifically, our results indicate that the instability driving the onset of locomotion is strengthened by substrate adhesiveness and weakened by cortical stiffness. Furthermore, the Fourier spectral analysis of morphology revealed lobe number n = 2 as the consistent dominant mode number across various experimental manipulations, suggesting that the instability mechanism driving the onset of Physarum locomotion is robust with respect to changes in environmental conditions and microplasmodial properties.