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Vol. 10, Issue 11, 3745-3769, November 1999
and
Center for Light Microscope Imaging and Biotechnology, Carnegie
Mellon University, Pittsburgh, Pennsylvania 15213
Forces generated by goldfish keratocytes and Swiss 3T3
fibroblasts have been measured with nanonewton precision and
submicrometer spatial resolution. Differential interference contrast
microscopy was used to visualize deformations produced by traction
forces in elastic substrata, and interference reflection microscopy
revealed sites of cell-substratum adhesions. Force ranged from a few
nanonewtons at submicrometer spots under the lamellipodium to several
hundred nanonewtons under the cell body. As cells moved forward,
centripetal forces were applied by lamellipodia at sites that remained
stationary on the substratum. Force increased and abruptly became
lateral at the boundary of the lamellipodium and the cell body. When
the cell retracted at its posterior margin, cell-substratum contact area decreased more rapidly than force, so that stress (force divided
by area) increased as the cell pulled away. An increase in lateral
force was associated with widening of the cell body. These mechanical
data suggest an integrated, two-phase mechanism of cell motility: (1)
low forces in the lamellipodium are applied in the direction of
cortical flow and cause the cell body to be pulled forward; and (2) a
component of force at the flanks pulls the rear margins forward toward
the advancing cell body, whereas a large lateral component contributes
to detachment of adhesions without greatly perturbing forward movement.
Online version of
this article contains video material for Figures 2, 4, 5, 6, 7,
9, and 10. Online version available at www.molbiolcell.org.
Present address: 11400 NE 132nd Street, L202,
Kirkland, WA 98034.
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