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Cells can sense forces applied to them, but also the stiffness of theirenvironment. These are two different phenomena, and here we investigate themechanosensitivity of the IInd kind: how the cell can measure an elasticmodulus at a single point of adhesion -- and how the cell can receive andinterpret the chemical signal released from the sensor. Our model uses theexample of large latent complex of TGFb as a sensor. Stochastic theory givesthe rate of breaking of latent complex, which initiates the signaling feedbackloop after the active TGFb release and leads to a change of cell phenotypedriven by the smooth muscle actin. We investigate the dynamic and steady-statebehaviours of the model, comparing them with experiments. In particular, weanalyse the timescale of approach to the steady state, the stability of thenon-linear dynamical system, and how the steady-state concentrations of the keymarkers vary depending on the elasticity of the substrate. We discover acrossover region for values of substrate elasticity closely corresponding tothat of the fibroblast to myofibroblast transition. We suggest that the cellcould actively vary the parameters of its dynamic feedback loop to `choose' theposition of the transition region and the range of substrate elasticity that itcan detect. In this way, the theory offers the unifying mechanism for a varietyof phenomena, such as the myofibroblast conversion in fibrosis of wounds andlungs and smooth muscle cell dysfunction in cardiac disease.

Mechanosensitivity of the IInd kind: TGFb mechanism of cell sensing the substrate stiffness