| Velocity transients and viscoelastic resistance to active shortening in cat papillary muscle Biophysical Journal, Volume 40, Issue 2, 1 November 1982, Pages 121-128 Y.L. Chiu, E.W. Ballou and L.E. Ford Abstract When isotonic force steps were applied to activated papillary muscles, the velocity was almost never constant. Early rapid shortening associated with the step persisted for 2–7 ms after the step ends. The early rapid shortening is attributed to lightly damped series elastic recoil and velocity transients of the contractile elements. In most steps, the subsequent velocity declines progressively with shortening, and most of the decline in velocity can be accounted for by compression of a viscoelastic element in parallel with the contractile elements. To demonstrate this, the time course of isotonic velocity was compared with a model in which the force-velocity characteristics of the contractile element were assumed to be constant, and the decline in velocity was due to increasing compression of the viscoelastic element. This model predicted the observed results except that the predicted velocities rose progressively above the measured values for steps to light loads applied late in the twitch, and fell below the velocity trace for heavy loads applied early in the twitch. These deviations would occur if rapid shortening caused deactivation late in the twitch, and if activation were rising early in the twitch. A conditioning step applied to the muscle during the rise of force depressed both isometric force and maximum velocity measured at the peak of force; isometric force was more depressed with later conditioning steps than with earlier steps, while maximum velocity was depressed by about the same extent with both early and late steps. This difference between the effects on isometric force and maximum velocity are explained by a combination of deactivation and viscoelastic load. Abstract | PDF (1145 kb) |
| Kinetic Effects of Fiber Type on the Two Subcomponents of the Huxley-Simmons Phase 2 in Muscle Biophysical Journal, Volume 85, Issue 1, 1 July 2003, Pages 390-401 Julien S. Davis and Neal D. Epstein Abstract The Huxley-Simmons phase 2 controls the kinetics of the first stages of tension recovery after a step-change in fiber length and is considered intimately associated with tension generation. It had been shown that phase 2 is comprised of two distinct unrelated phases. This is confirmed here by showing that the properties of phase 2 are independent of fiber type, whereas those of phase 2 are fiber type dependent. Phase 2 has a rate of 1000–2000s and is temperature insensitive (∼1.16) in fast, medium, and slow speed fibers. Regardless of fiber type and temperature, the amplitude of phase 2 is half (∼0.46) that of phase 1 (fiber instantaneous stiffness). Consequently, fiber compliance (cross-bridge and thick/thin filament) appears to be the common source of both phase 1 elasticity and phase 2 viscoelasticity. In fast fibers, stiffness increases in direct proportion to tension from an extrapolated positive origin at zero tension. The simplest explanation is that tension generation can be approximated by two-state transition from attached preforce generating (moderate stiffness) to attached force generating (high stiffness) states. Phase 2 is quite different, progressively slowing in concert with fiber type. An interesting interpretation of the amplitude and rate data is that reverse coupling of phase 2 back to P release and ATP hydrolysis appears absent in fast fibers, detectable in medium speed fibers, and marked in slow fibers contracting isometrically. Contracting slow and heart muscles stretched under load could employ this enhanced reversibility of the cross-bridge cycle as a mechanism to conserve energy. Abstract | Full Text | PDF (176 kb) |
| Viscoelastic Retraction of Single Living Stress Fibers and Its Impact on Cell Shape, Cytoskeletal Organization, and Extracellular Matrix Mechanics Biophysical Journal, Volume 90, Issue 10, 15 May 2006, Pages 3762-3773 Sanjay Kumar, Iva Z. Maxwell, Alexander Heisterkamp, Thomas R. Polte, Tanmay P. Lele, Matthew Salanga, Eric Mazur and Donald E. Ingber Abstract Cells change their form and function by assembling actin stress fibers at their base and exerting traction forces on their extracellular matrix (ECM) adhesions. Individual stress fibers are thought to be actively tensed by the action of actomyosin motors and to function as elastic cables that structurally reinforce the basal portion of the cytoskeleton; however, these principles have not been directly tested in living cells, and their significance for overall cell shape control is poorly understood. Here we combine a laser nanoscissor, traction force microscopy, and fluorescence photobleaching methods to confirm that stress fibers in living cells behave as viscoelastic cables that are tensed through the action of actomyosin motors, to quantify their retraction kinetics in situ, and to explore their contribution to overall mechanical stability of the cell and interconnected ECM. These studies reveal that viscoelastic recoil of individual stress fibers after laser severing is partially slowed by inhibition of Rho-associated kinase and virtually abolished by direct inhibition of myosin light chain kinase. Importantly, cells cultured on stiff ECM substrates can tolerate disruption of multiple stress fibers with negligible overall change in cell shape, whereas disruption of a single stress fiber in cells anchored to compliant ECM substrates compromises the entire cellular force balance, induces cytoskeletal rearrangements, and produces ECM retraction many microns away from the site of incision; this results in large-scale changes of cell shape (> 5% elongation). In addition to revealing fundamental insight into the mechanical properties and cell shape contributions of individual stress fibers and confirming that the ECM is effectively a physical extension of the cell and cytoskeleton, the technologies described here offer a novel approach to spatially map the cytoskeletal mechanics of living cells on the nanoscale. Abstract | Full Text | PDF (396 kb) |
Copyright © 1970 The Biophysical Society. All rights reserved.
Biophysical Journal, Volume 10, Issue 6, 539-555, 1 June 1970
doi:10.1016/S0006-3495(70)86318-4
Articles
Julia T. Apter and William W. Graessley
A model for muscular behavior has been developed by a generalization of the laws governing the viscoelastic behavior of polymeric materials. The model simulates events thought to take place during stretch, loading, and stimulation of muscle, whether smooth or striated. The equations of motion were solved with an analogue computer for several types of perturbation, and stress, strain, and strainrate curves were generated. Model parameters were selected by fitting experimental stress-relaxation data. The resulting equations predicted the frequency dependence of dynamic modulus and phase angle within experimental error. With appropriate boundary conditions and suitable values for model parameters, the computed results also closely resembled experimental curves of contraction velocity vs. time, isometric tension development vs. time, force-velocity curves, and temperature-tension relationships. These results call attention to the relationship between the behavior of various kinds of muscle and open the way for quantifying muscular behavior in general.