We used a novel uniaxial stretching rheometer to measure the creep function J(t) of an isolated living cell. We show, for the first time at the scale of the whole cell, that J(t) behaves as a power-law J(t) = At(alpha). For N = 43 mice myoblasts (C2-7), we find alpha = 0.24 +/- 0.01 and A = (2.4 +/- 0.3) 10(-3) Pa(-1) s(-alpha). Using Laplace Transforms, we compare A and alpha to the parameters G(0) and beta of the complex modulus G*(omega) = G(0)omega(beta) measured by other authors using magnetic twisting cytometry and atomic force microscopy. Excellent agreement between A and G(0) on the one hand, and between alpha and beta on the other hand, indicated that the power-law is an intrinsic feature of cell mechanics and not the signature of a particular technique. Moreover, the agreement between measurements at very different size scales, going from a few tens of nanometers to the scale of the whole cell, suggests that self-similarity could be a central feature of cell mechanical structure. Finally, we show that the power-law behavior could explain previous results first interpreted as instantaneous elasticity. Thus, we think that the living cell must definitely be thought of as a material with a large and continuous distribution of relaxation time constants which cannot be described by models with a finite number of springs and dash-pots.
The area expansion and the shear moduli of the free spectrin skeleton, freshly extracted from the membrane of a human red blood cell (RBC), are measured by using optical tweezers micromanipulation. An RBC is trapped by three silica beads bound to its membrane. After extraction, the skeleton is deformed by applying calibrated forces to the beads. The area expansion modulus K(C) and shear modulus mu(C) of the two-dimensional spectrin network are inferred from the deformations measured as functions of the applied stress. In low hypotonic buffer (25 mOsm/kg), one finds K(C) = 4.8 +/- 2.7 microN/m, mu(C) = 2.4 +/- 0.7 microN/m, and K(C)/mu(C) = 1.9 +/- 1.0. In isotonic buffer, one measures higher values for K(C), mu(C), and K(C)/mu(C), partly because the skeleton collapses in a high-ionic-strength environment. Some data concerning the time evolution of the mechanical properties of the skeleton after extraction and the influence of ATP are also reported. In the Discussion, it is shown that the measured values are consistent with estimates deduced from experiments carried out on the intact membrane and agree with theoretical and numerical predictions concerning two-dimensional networks of entropic springs.
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