Epithelial cells fluidize upon adhesion but display mechanical homeostasis in the adherent state

Atomic force microscopy is used to study the viscoelastic properties of epithelial cells in three different states. Force relaxation data are acquired from cells in suspension, adhered but single cells, and polarized cells in a confluent monolayer using different indenter geometries comprising flat bars, pyramidal cones, and spheres. We found that the fluidity of cells increased substantially from the suspended to the adherent state. Along this line, the prestress of suspended cells generated by cortical contractility is also greater than that of cells adhering to a surface. Polarized cells that are part of a confluent monolayer form an apical cap that is soft and fluid enough to respond rapidly to mechanical challenges from wounding, changes in the extracellular matrix, osmotic stress, and external deformation. In contrast to adherent cells, cells in the suspended state show a pronounced dependence of fluidity on the external areal strain. With increasing areal strain, the suspended cells become softer and more fluid. We interpret the results in terms of cytoskeletal remodeling that softens cells in the adherent state to facilitate adhesion and spreading by relieving internal active stress. However, once the cells spread on the surface they maintain their mechanical phenotype displaying viscoelastic homeostasis.