Fig. 8. A schematic model of how the cell exerts the traction force against the
substrate during migration. The flexible silicone substrate is shown as
straight or wavy lines. The wavy lines indicate the distortion of the silicone
surface. The gray large arrows (a,f) show the direction of the cell movement.
The circles and crosses indicate the present and the original position of
beads, respectively. The displacement of beads is shown by thin black arrows
parallel to the substrate. The shaded part of the cell represents the putative
region generating motive force for migration. The cell transmits the traction
force or the motive force (thick black arrows) to the substrate through
putative feet (black vertical bars). (a-e) In the wild-type cell, during the
extension phase (a-c), the extension force of the anterior region drags the
posterior cell body and causes the pulling traction force at the anterior
feet, and the posterior feet are passively dragged and pull the beads forward.
During this phase, formation of new feet may occur in the newly extended
anterior region (c,d). During the retraction phase (d,e), the posterior region
of the cell contracts (two small arrows at the rear) mediated by the myosin
II, which localizes at this region. It causes the detachment of the adhesion
sites in the posterior region, the substrate is relaxed, and the beads return
to their original positions (the circle overlaps with the cross) in d. (e)
Then the posterior contraction generates a pushing force of the anterior cell
body and also causes anterior extension. At this time, the posterior feet
exert the traction force backward to the substrate and the anterior feet push
the beads forward. (f-i) In the case of MHC-null cells, the beads always move
toward the cell body since the contraction of posterior regions is deficient.
The posterior edge moves at a constant rate whereas the extension progresses
cyclically at the anterior region. Figs j and k explain the difference between
the `pulling force' and the `pushing force' at the anterior region. (j) The
pulling force is caused by the reaction to the anterior motive force generated
by actin polymerization or myosin Is. (k) The pushing force is generated by
the posterior contraction mediated by myosin II.