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15.2 METHODOLOGY                                      293

                                                      FIG. 15.4  Schematic illustration of the cell deformation resulting from the sensing
                                                      force in the cell mechanosensing stage. N i is a point on the surface of the undeformed
                                                      cell (solid line), n i is the same point on the surface of the deformed cell (dashed line), and
                                                      C is the cell center.




















                                                      F trac  ¼ σ cell                                      (15.5)
                                                        i    i  Skn r ψe i
           where S stands for a proportionality model parameter with units of area. k is the binding constant for the integrins at
           the front and rear of the cell to the ligands in the ECM and n r is the total number of available receptors at the front and
           rear of the cell. Finally, ψ represents the concentration of the ligands at the leading edge of the cell in the ECM [27].
                                              net
           Therefore, The resultant traction force, F trac , can be calculated by the summation of the traction forces applied at each
           node as
                                                               n
                                                              X
                                                        F net  ¼  F trac                                    (15.6)
                                                                  i
                                                         trac
                                                              i¼1
           where n is the number of the nodes located on the cell surface.
              In a multicellular system, cells deform when they contact each other to occupy all the ECM [72, 100]. Therefore,
           when cell migration is considered with a constant cell shape, to avoid interference of two cells it is assumed that
                                                         k r j  r i k  2r                                   (15.7)
           where x i and x j are the position vector of each cell centroid (Fig. 15.5).
              During the mechanosensing process, when two or more cells come into contact with each other, their in-common
           nodal points on the cell surface are not able to send out any pseudopods to sense the ECM [100, 103, 104]. Therefore,
           these cells do not exert any sensing force at those nodes until they are separated again. However, in these nodes, the


                                                   FIG. 15.5  Two in-contact cells with four shared nodes, n 1 , n 2 , n 3 , and n 4 .



























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