Asymmetry between pushing and pulling for crawling cells.

Eukaryotic cells possess motility mechanisms allowing them not only to self-propel but also to exert forces on obstacles (to push) and to carry cargoes (to pull). To study the inherent asymmetry between active pushing and pulling we model a crawling acto-myosin cell extract as a one-dimensional layer of active gel subjected to external forces. We show that pushing is controlled by protrusion and that the macroscopic signature of the protrusion dominated motility mechanism is concavity of the force-velocity relation. In contrast, pulling is driven by protrusion only at small values of the pulling force and it is replaced by contraction when the pulling force is sufficiently large. This leads to more complex convex-concave structure of the force-velocity relation; in particular, competition between protrusion and contraction can produce negative mobility in a biologically relevant range. The model illustrates active readjustment of the force generating machinery in response to changes in the dipole structure of external forces. The possibility of switching between complementary active mechanisms implies that if necessary "pushers" can replace "pullers" and vice versa.