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Isostaticity and Controlled Force Transmission in the Cytoskeleton: A Model Awaiting Experimental Evidence

Biological and Soft Systems, Cavendish Laboratory, Cambridge, United Kingdom; and Earth Science and Engineering, Imperial College, London, United Kingdom

Correspondence: Address reprint requests to Raphael Blumenfeld, Biological and Soft Systems, Cavendish Laboratory, Madingley Road, Cambridge CB3 0HE, UK. E-mail: rbb11{at}phy.cam.ac.uk.

A new model is proposed for force transmission through the cytoskeleton (CSK). A general discussion is first presented on the physical principles that underlie the modeling of this phenomenon. Some fundamental problems of conventional models—continuous and discrete—are examined. It is argued that mediation of focused forces is essential for good control over intracellular mechanical signals. The difficulties of conventional continuous models in describing such mediation are traced to a fundamental assumption rather than to their being continuous. Relevant advantages and disadvantages of continuous and discrete modeling are discussed. It is concluded that favoring discrete models is based on two misconceptions, which are clarified. The model proposed here is based on the idea that focused propagation of mechanical stimuli in frameworks over large distances (compared to the mesh size) can only occur when considerable regions of the CSK are isostatic. The concept of isostaticity is explained and a recently developed continuous isostaticity theory is briefly reviewed. The model enjoys several advantages: it leads to good control over force mediation; it explains nonuniform stresses and action at a distance; it is continuous, making it possible to model force propagation over long distances; and it enables prediction of individual force paths. To be isostatic, or nearly so, CSK networks must possess specific structural characteristics, and these are quantified. Finally, several experimental observations are interpreted using the new model and implications are discussed. It is also suggested that this approach may give insight into the dynamics of reorganization of the CSK. Many of the results are amenable to experimental measurements, providing a testing ground for the proposed picture, and generic experiments are suggested.