Mechanics of crawling cells.

Crawling of keratocytes derived from aquatic vertebrates represents a very useful model system for the investigation of cell locomotion because of its ease of handling and the clear structural separation of a thin cytoplasmic layer, the lamella, from the cell body containing the nucleus and other organelles. Spreading of spherical keratocytes results in fried egg shaped cells, which on withdrawing their lamella at one side become polarized and start moving. Hydrostatic pressure, tension at the cortex, traction forces exerted on the adhesion sites and inside the cells along filamentous structures are required to gain a certain shape. Traction forces have been made visible using scanning acoustic microscopy. This method also allowed for the demonstration of cytoplasmic fluxes inside a moving keratocyte and changes of forces while a migrating cell is changing its direction of locomotion. The pros and cons for actin polymerization at the leading front providing the driving force for crawling are discussed on the basis of structural and experimental results: do they stringently identify polymerization of actin as the only driving machinery. Such a mechanism not only should explain the advancement of the leading edge but also the movement of the whole cell, i.e. the material flux taking place from the cell body to the periphery. Even if the lamella periphery itself may be motile by actin turnover this scheme may represent an oversimplification if applied to the whole cell. Considering the complexity of a whole cell simplifying model systems may not lead to adequate descriptions of the mechanisms as they occur within cells with a highly complex structure, although the model might be consistent and sufficient to describe, i.e. crawling in general.