“…Those forces come from the cortical cytoskeleton pulling against the deeper cytoskeleton and the substrates. If the cortex should separate from the deeper cytoskeleton, as it does in blebs ( Charras et al, 2005 ; Moeendarbary et al, 2013 ), the cortex becomes stiff because it is under a hydrostatic pressure gradient ( Beyder and Sachs, 2009 , 2011 ). We found that in HEK cells, about half of the cortical stress is in the bilayer and half in the attached proteins ( Akinlaja and Sachs, 1998 ).…”