arXiv subjects

Lev Truskinovsky

Publications and source records attributed to Lev Truskinovsky.

At least 37 records · Page 2Linked to original sources

arXiv2021

A plausible mechanism of muscle stabilization in stall conditions

We address the well-known limitation of the Huxley and Simmons 1971 (HS) model. It is a statement that at physiological value of stiffness in the actomyosin complex, the distribution of the myosin motors becomes microscopically uniform (all the motors are either in pre- or post-power stroke conformation) after an infinitesimal displacement from the stall (isometric contractions) conditions. Such uniform behavior at the fiber level would generate a negative slope in the $T_2-δ$ relationship (in the nomenclature of the HS paper), not observed experimentally. This negative slope means inhomogeneity of the macroscopic sarcomere configuration, which is also not observed. To address this controversial prediction of the HS theory, we explore the possibility that the slope of the $T_2-δ$ curve is, in fact, positive due to an interaction between neighboring cross-bridges. We show that such interaction can potentially destabilize the uniform configurations (all pre or all post) by making the non-uniform configurations energetically preferable. We argue that, despite the presence of other factors, which can in principle also ensure the microscopic inhomogeneity of cross-bridge configurations, the implied interaction is an important player in muscle mechanics.

physics.bio-ph↗

arXiv2021

Fluctuations in crystalline plasticity

Recently acoustic signature of dislocation avalanches in HCP materials was found to be long tailed in size and energy, suggesting critical dynamics. Even more recently, the intermittent plastic response was found to be generic for micro- and nano-sized systems independently of their crystallographic symmetry. These rather remarkable discoveries are reviewed in this paper in the perspective of the recent studies performed in our group. We discuss the physical origin and the scaling properties of plastic fluctuations and address the nature of their dependence on crystalline symmetry, system size, and disorder content. A particular emphasis is placed on the associated emergent behaviors, including the formation of dislocation structures, and on our ability to temper plastic fluctuations by alloying. We also discuss the "smaller is wilder" size effect that culminates in a paradoxical crack-free brittle behavior of very small, initially dislocation free crystals. We show that the implied transition between different rheological behaviors is regulated by the ratio of length scales $R=L/l$, where $L$ is the system size and $l$ is the internal length. We link this new size effect with other related phenomena like size dependence of strength ("smaller is stronger") and the size induced switch between different hardening mechanisms. One of the technological challenges in nanoscience is to tame the intermittency of plastic flow. We show that this task can be accomplished by generating tailored quenched disorder which allows one to control micro- and nano-scale forming and opens new perspectives in micro-metallurgy and structural engineering of ultra-small load-carrying elements. These results could not be achieved by conventional methods that do not explicitly consider the stochastic nature of collective dislocation dynamics.

cond-mat.mes-hall↗

arXiv2020

Peristalsis by pulses of activity

Peristalsis by actively generated waves of muscle contraction is one of the most fundamental ways of producing motion in living systems. We show that peristalsis can be modeled by a train of rectangular-shaped solitary waves of localized activity propagating through otherwise passive matter. Our analysis is based on the FPU-type discrete model accounting for active stresses and we reveal the existence in this problem of a critical regime which we argue to be physiologically advantageous.

physics.bio-ph↗

arXiv2020

Rigidity-Controlled Crossover: From Spinodal to Critical Failure

Failure in disordered solids is accompanied by intermittent fluctuations extending over a broad range of scales. The implied scaling has been previously associated with either spinodal or critical points. We use an analytically transparent mean-field model to show that both analogies are relevant near the brittle-to-ductile transition. Our study indicates that in addition to the strength of quenched disorder, an appropriately chosen global measure of rigidity (connectivity) can be also used to tune the system to criticality. By interpreting rigidity as a timelike variable we reveal an intriguing parallel between earthquake-type critical failure and Burgers turbulence.

cond-mat.dis-nn↗

arXiv2019

Functionality of disorder in muscle mechanics

A salient feature of skeletal muscles is their ability to take up an applied slack in a microsecond timescale. Behind this remarkably fast adaptation is a collective folding in a bundle of elastically interacting bistable elements. Since this interaction has long-range character, the behavior of the system in force and length controlled ensembles is different; in particular, it can have two distinct order-disorder--type critical points. We show that the account of the disregistry between myosin and actin filaments places the elementary force-producing units of skeletal muscles close to both such critical points. The ensuing "double-criticality" contributes to the system's ability to perform robustly and suggests that the disregistry is functional.

physics.bio-ph↗

arXiv2019

Supersonic kinks and solitons in active solids

To show that steadily propagating nonlinear waves in active matter can be driven internally, we develop a prototypical model of a topological kink moving with a constant supersonic speed. We use a model of a bi-stable mass-spring (FPU) chain capable of generating active stress. In contrast to subsonic kinks in passive bi-stable chains, that are necessarily dissipative, the obtained supersonic solutions are purely anti-dissipative. Our numerical experiments point towards stability of the obtained kink-type solutions and the possibility of propagating kink-anti-kink bundles reminiscent of solitons. We show that even the simplest quasi-continuum approximation of the discrete model captures the most important features of the predicted active phenomena.

cond-mat.soft↗

arXiv2019

Passive viscoelastic response of striated muscles

Muscle cells with sarcomeric structure exhibit highly nontrivial passive mechanical response. The difficulty of its continuum modeling is due to the presence of long-range interactions transmitted by extended protein skeleton. To build a rheological model for muscle 'material' we use a stochastic micromodel and derive a linear response theory for a half-sarcomere. Instead of the first order rheological equation, anticipated by A.V. Hill on the phenomenological grounds, we obtain a novel second order equation. We use the values of the microscopic parameters for frog muscles to show that the proposed rheological model is in excellent quantitative agreement with physiological experiments.

physics.bio-ph↗

arXiv2019

Snare machinery is optimized for ultrafast fusion

SNARE proteins zipper to form SNAREpins that power vesicle fusion with target membranes in a variety of biological processes. A single SNAREpin takes about 1 second to fuse two bilayers, yet a handful can ensure release of neurotransmitters from synaptic vesicles much faster, in a 10th of a millisecond. We propose that, similar to the case of muscle myosins, the ultrafast fusion results from cooperative action of many SNAREpins. The coupling originates from mechanical interactions induced by confining scaffolds. Each SNAREpin is known to have enough energy to overcome the fusion barrier of 25-35 kB T, however, the fusion barrier only becomes relevant when the SNAREpins are nearly completely zippered and from this state each SNAREpin can deliver only a small fraction of this energy as mechanical work. Therefore they have to act cooperatively and we show that at least 3 of them are needed to ensure fusion in less than a millisecond. However, to reach the pre-fusion state collectively, starting from the experimentally observed half-zippered metastable state, the SNAREpins have to mechanically synchronize which takes exponentially longer time as the number of SNAREpins increases. Incorporating this somewhat counter-intuitive idea in a simple coarse grained model results in the novel prediction that there should be an optimum number of SNAREpins for sub-ms fusion: 3-6 over a wide range of parameters. Interestingly, in situ cryo-electron microscope tomography has very recently shown that exactly six SNAREpins participate in the fusion of each synaptic vesicle. This number is in the range predicted by our theory.

physics.bio-ph↗

arXiv2019

Nonlinear elasticity of incompatible surface growth

Surface growth is a crucial component of many natural and artificial processes from cell proliferation to additive manufacturing. In elastic systems surface growth is usually accompanied by the development of geometrical incompatibility leading to residual stresses and triggering various instabilities. In a recent paper (PRL, 119, 048001, 2017) we developed a linearized elasticity theory of incompatible surface growth which quantitatively linked deposition protocols with post-growth states of stress. Here we extend this analysis to account for both physical and geometrical nonlinearities of an elastic solid. The new development reveals the shortcomings of the linearized theory, in particular, its inability to describe kinematically confined surface growth and to account for growth-induced elastic instabilities.

cond-mat.soft↗

arXiv2018

Drift induced by dissipation

Active particles have become a subject of intense interest across several disciplines from animal behavior to granular physics. Usually the models of such particles contain an explicit internal driving. Here we propose a model with implicit driving in the sense that the behavior of our particle is fully dissipative at zero temperature but becomes active in the presence of seemingly innocent equilibrium fluctuations. The mechanism of activity is related to the breaking of the gradient structure in the chemo-mechanical coupling. We show that the thermodynamics of such active particles depends crucially on inertia and cannot be correctly captured in the standard Smoluchowski limit. To deal with stall conditions, we generalize the definition of Stokes efficiency, assessing the quality of active force generation. We propose a simple realization of the model in terms of an electric circuit capable of turning fluctuations into a directed current without an explicit source of voltage.

cond-mat.stat-mech↗

arXiv2018

Colliding cells: when active segments behave as active particles

Quantifying the outcomes of cells collisions is a crucial step in building the foundations of a kinetic theory of living matter. Here, we develop a mechanical theory of such collisions by first representing individual cells as extended objects with internal activity and then reducing this description to a model of size-less active particles characterized by their position and polarity. We show that, in the presence of an applied force, a cell can either be dragged along or self-propel against the force, depending on the polarity of the cell. The co-existence of these regimes offers a self-consistent mechanical explanation for cell re-polarization upon contact. We rationalize the experimentally observed collision scenarios within the extended and particle models and link the various outcomes with measurable biological parameters.

physics.bio-ph↗

arXiv2018

Physics of muscle contraction

In this paper we report, clarify and broaden various recent efforts to complement the chemistry-centered models of force generation in muscles by mechanics-centered models. The physical mechanisms of interest can be grouped into two classes: passive and active. The main passive effect is the fast force recovery which does not require the detachment of myosin cross-bridges from actin filaments and can operate without a specialized supply ATP. In mechanical terms, it can be viewed as a collective folding-unfolding phenomenon in the system of interacting bistable units and modeled by near equilibrium Langevin dynamics. The parallel active force generation mechanism operates at slow timescales, requires detachment and is crucially dependent on ATP hydrolysis. The underlying mechanical processes take place far from equilibrium and are represented by stochastic models with broken time reversal symmetry implying non-potentiality, correlated noise or multiple reservoirs. The modeling approaches reviewed in this paper deal with both active and passive processes and support from the mechanical perspective the biological point of view that phenomena involved in slow (active) and fast (passive) force generation are tightly intertwined. They reveal, however, that biochemical studies in solution, macroscopic physiological measurements and structural analysis do not provide by themselves all the necessary insights into the functioning of the contractile system. In particular, the reviewed body of work emphasizes the important role of long-range interactions and criticality in securing the targeted mechanical response in the physiological regime of isometric contractions. The importance of the purely mechanical microscale modeling is accentuated at the end of the paper where we address the puzzling issue of the stability of muscle response on the so called descending limb of the isometric tetanus.

physics.bio-ph↗

arXiv2018

Mechanical stress as a regulator of cell motility

The motility of a cell can be triggered or inhibited not only by an applied force but also by a mechanically neutral force couple. This type of loading, represented by an applied stress and commonly interpreted as either squeezing or stretching, can originate from extrinsic interaction of a cell with its neighbors. To quantify the effect of applied stresses on cell motility we use an analytically transparent one-dimensional model accounting for active myosin contraction and induced actin turnover. We show that stretching can polarize static cells and initiate cell motility while squeezing can symmetrize and arrest moving cells. We show further that sufficiently strong squeezing can lead to the loss of cell integrity. The overall behavior of the system depends on the two dimensionless parameters characterizing internal driving (chemical activity) and external loading (applied stress). We construct a phase diagram in this parameter space distinguishing between, static, motile and collapsed states. The obtained results are relevant for the mechanical understanding of contact inhibition and the epithelial-to-mesenchymal transition.

cond-mat.soft↗

arXiv2017

Inelastic Surface Growth

Inelastic surface growth associated with continuous creation of incompatibility on the boundary of an evolving body is behind a variety of natural and technological processes, including embryonic development and 3D printing. In this paper we extend the recently proposed stress-space-centered theory of surface growth (PRL 119, 048001, 2017) by shifting the focus towards growth induced strains. To illustrate the new development we present several analytically tractable examples.

cond-mat.soft↗

arXiv2017

Cooperative folding of muscle myosins: I. Mechanical model

Mechanically induced folding of passive cross-linkers is a fundamental biological phenomenon. A typical example is a conformational change in myosin II responsible for the power-stroke in skeletal muscles. In this paper we present an athermal perspective on such folding by analyzing the simplest purely mechanical prototype: a parallel bundle of bi-stable units attached to a common backbone. We show that in this analytically transparent model, characterized by a rugged energy landscape, the ground states are always highly coherent, single-phase configurations. We argue that such cooperative behavior, ensuring collective conformational change, is due to the dominance of long- range interactions making the system non-additive. The detailed predictions of our model are in agreement with experimentally observed non-equivalence of fast force recovery in skeletal muscles loaded in soft and hard devices. Some features displayed by the model are also recognizable in the behavior of other biological systems with passive multi-stability and long-range interactions including detaching adhesive binders and pulled RNA/DNA hairpins.

physics.bio-ph↗

arXiv2017

Paramagnetic response of muscle-type systems

We provide a prototypical description of mechanically-induced collective conformational change (folding), relevant in a variety of biological contexts from muscle contraction to hair cell gating and integrin binding. Our study is inspired by the seminal Huxley-Simmons (HS) model which we reinterpret from the perspective of stochastic dynamics involving rigidly coupled bi-stable elements. In this interpretation the HS model can be mapped to a paramagnetic Ising model, however, the equivalence is not complete due to the presence of elastic elements responsible for negative susceptibility and quasi-critical behavior. We go beyond the conventional chemo-mechanical description of such systems, revealing both its strengths and its limitations.

physics.bio-ph↗

arXiv2017

Bi-stability resistant to fluctuations

We study a simple micro-mechanical device that does not lose its snap-through behavior in an environment dominated by fluctuations. The main idea is to have several degrees of freedom that can cooperatively resist the de-synchronizing effect of random perturbations. As an inspiration we use the power stroke machinery of skeletal muscles, which ensures at sub-micron scales and finite temperatures a swift recovery of an abruptly applied slack. In addition to hypersensitive response at finite temperatures, our prototypical Brownian snap spring also exhibits criticality at special values of parameters which is another potentially interesting property for micro-scale engineering applications.

physics.bio-ph↗

arXiv2017

Printing non-Euclidean solids

Geometrically frustrated solids with non-Euclidean reference metric are ubiquitous in biology and are becoming increasingly relevant in technological applications. Often they acquire a targeted con- figuration of incompatibility through surface accretion of mass as in tree growth or dam construction. We use the mechanics of incompatible surface growth to show that geometrical frustration develop- ing during deposition can be fine-tuned to ensure a particular behavior of the system in physiological (or working) conditions. As an illustration, we obtain an explicit 3D printing protocol for arteries, which guarantees stress uniformity under inhomogeneous loading, and for explosive plants, allowing a complete release of residual elastic energy with a single cut. Interestingly, in both cases reaching the physiological target requires the incompatibility to have a topological (global) component.

cond-mat.soft↗