Mechanics and force transmission in soft composites of rods in elastic gels

We report detailed theoretical investigations of the micro-mechanics and bulk elastic properties of composites consisting of randomly distributed stiff fibers embedded in an elastic matrix in two and three dimensions. Recent experiments published in Physical Review Letters [102, 188303 (2009)] have suggested that the inclusion of stiff microtubules in a softer, nearly incompressible biopolymer matrix can lead to emergent compressibility. This can be understood in terms of the enhancement of the compressibility of the composite relative to its shear compliance as a result of the addition of stiff rod-like inclusions. We show that the Poisson's ratio $\nu$ of such a composite evolves with increasing rod density towards a particular value, or {\em fixed point}, independent of the material properties of the matrix, so long as it has a finite initial compressibility. This fixed point is $\nu=1/4$ in three dimensions and $\nu=1/3$ in two dimensions. Our results suggest an important role for stiff filaments such as microtubules and stress fibers in cell mechanics. At the same time, our work has a wider elasticity context, with potential applications to composite elastic media with a wide separation of scales in stiffness of its constituents such as carbon nanotube-polymer composites, which have been shown to have highly tunable mechanics.Comment: 10 pages, 8 figure

Effect of dipolar moments in domain sizes of lipid bilayers and monolayers

Lipid domains are found in systems such as multi-component bilayer membranes and single component monolayers at the air-water interface. It was shown by Andelman et al. (Comptes Rendus 301, 675 (1985)) and McConnell et al. (Phys. Chem. {\bf 91}, 6417 (1987)) that in monolayers, the size of the domains results from balancing the line tension, which favors the formation of a large single circular domain, against the electrostatic cost of assembling the dipolar moments of the lipids. In this paper, we present an exact analytical expression for the electric potential, ion distribution and electrostatic free energy for different problems consisting of three different slabs with different dielectric constants and Debye lengths, with a circular homogeneous dipolar density in the middle slab. From these solutions, we extend the calculation of domain sizes for monolayers to include the effects of finite ionic strength, dielectric discontinuities (or image charges) and the polarizability of the dipoles and further generalize the calculations to account for domains in lipid bilayers. In monolayers, the size of the domains is dependent on the different dielectric constants but independent of ionic strength. In asymmetric bilayers, where the inner and outer leaflets have different dipolar densities, domains show a strong size dependence with ionic strength, with molecular-sized domains that grow to macroscopic phase separation with increasing ionic strength. We discuss the implications of the results for experiments and briefly consider their relation to other two dimensional systems such as Wigner crystals or heteroepitaxial growth.Comment: 13 pages, 5 figues in eps Replaced with new version, one citation added and a few statements corrected. The results of the paper are unchange

Non-local fluctuation correlations in active gels

Many active materials and biological systems are driven far from equilibrium by embedded agents that spontaneously generate forces and distort the surrounding material. Probing and characterizing these athermal fluctuations is essential for understanding the properties and behaviors of such systems. Here we present a mathematical procedure to estimate the local action of force-generating agents from the observed fluctuating displacement fields. The active agents are modeled as oriented force dipoles or isotropic compression foci, and the matrix on which they act is assumed to be either a compressible elastic continuum or a coupled network-solvent system. Correlations at a single point and between points separated by an arbitrary distance are obtained, giving a total of three independent fluctuation modes that can be tested with microrheology experiments. Since oriented dipoles and isotropic compression foci give different contributions to these fluctuation modes, ratiometric analysis allows us characterize the force generators. We also predict and experimentally find a high-frequency ballistic regime, arising from individual force generating events in the form of the slow build-up of stress followed by rapid but finite decay. Finally, we provide a quantitative statistical model to estimate the mean filament tension from these athermal fluctuations, which leads to stiffening of active networks.Comment: 12 pages, 7 figures; some clarifications and ammended figure notation

A core genetic module : the Mixed Feedback Loop

The so-called Mixed Feedback Loop (MFL) is a small two-gene network where protein A regulates the transcription of protein B and the two proteins form a heterodimer. It has been found to be statistically over-represented in statistical analyses of gene and protein interaction databases and to lie at the core of several computer-generated genetic networks. Here, we propose and mathematically study a model of the MFL and show that, by itself, it can serve both as a bistable switch and as a clock (an oscillator) depending on kinetic parameters. The MFL phase diagram as well as a detailed description of the nonlinear oscillation regime are presented and some biological examples are discussed. The results emphasize the role of protein interactions in the function of genetic modules and the usefulness of modelling RNA dynamics explicitly.Comment: To be published in Physical Review

Filamin cross-linked semiflexible networks: Fragility under strain

The semiflexible F-actin network of the cytoskeleton is cross-linked by a variety of proteins including filamin, which contain Ig-domains that unfold under applied tension. We examine a simple semiflexible network model cross-linked by such unfolding linkers that captures the main mechanical features of F-actin networks cross-linked by filamin proteins and show that under sufficiently high strain the network spontaneously self-organizes so that an appreciable fraction of the filamin cross-linkers are at the threshold of domain unfolding. We propose an explanation of this organization based on a mean-field model and suggest a qualitative experimental signature of this type of network reorganization under applied strain that may be observable in intracellular microrheology experiments of Crocker et al.Comment: 4 Pages, 3 figures, Revtex4, submitted to PR

Testing Tensile and Shear Epoxy Strength at Cryogenic Temperatures

This paper covers cryogenic, tensile testing and research completed on a number of epoxies used in cryogenic applications. Epoxies are used in many different applications; however, this research focused on the use of epoxy used to bond MLI standoffs to cryogenic storage tanks and the loads imparted to the tank through the MLI. To conduct testing, samples were made from bare stainless steel, aluminum and primed aluminum. Testing involved slowly cooling test samples with liquid nitrogen then applying gradually increasing tensile loads to the epoxy. The testing evaluated the strength and durability of epoxies at cryogenic temperatures and serves as a base for future testing. The results of the tests showed that some epoxies withstood the harsh conditions while others failed. The two epoxies yielding the best results were Masterbond EP29LPSP and Scotch Weld 2216. For all metal surfaces tested, both epoxies had zero failures for up to 11.81 kg of mass

A two-state model for helicase translocation and unwinding of nucleic acids

Helicases are molecular motors that unwind double-stranded nucleic acids (dsNA), such as DNA and RNA). Typically a helicase translocates along one of the NA single strands while unwinding and uses adenosine triphosphate (ATP) hydrolysis as an energy source. Here we model of a helicase motor that can switch between two states, which could represent two different points in the ATP hydrolysis cycle. Our model is an extension of the earlier Betterton-J\"ulicher model of helicases to incorporate switching between two states. The main predictions of the model are the speed of unwinding of the dsNA and fluctuations around the average unwinding velocity. Motivated by a recent claim that the NS3 helicase of Hepatitis C virus follows a flashing ratchet mechanism, we have compared the experimental results for the NS3 helicase with a special limit of our model which corresponds to the flashing ratchet scenario. Our model accounts for one key feature of the experimental data on NS3 helicase. However, contradictory observations in experiments carried out under different conditions limit the ability to compare the model to experiments.Comment: minor modification

Probing molecular free energy landscapes by periodic loading

Single molecule pulling experiments provide information about interactions in biomolecules that cannot be obtained by any other method. However, the reconstruction of the molecule's free energy profile from the experimental data is still a challenge, in particular for the unstable barrier regions. We propose a new method for obtaining the full profile by introducing a periodic ramp and using Jarzynski's identity for obtaining equilibrium quantities from non-equilibrium data. Our simulated experiments show that this method delivers significant more accurate data than previous methods, under the constraint of equal experimental effort.Comment: 4 pages, 3 figure

Proton transport and torque generation in rotary biomotors

We analyze the dynamics of rotary biomotors within a simple nano-electromechanical model, consisting of a stator part and a ring-shaped rotor having twelve proton-binding sites. This model is closely related to the membrane-embedded F$_0$ motor of adenosine triphosphate (ATP) synthase, which converts the energy of the transmembrane electrochemical gradient of protons into mechanical motion of the rotor. It is shown that the Coulomb coupling between the negative charge of the empty rotor site and the positive stator charge, located near the periplasmic proton-conducting channel (proton source), plays a dominant role in the torque-generating process. When approaching the source outlet, the rotor site has a proton energy level higher than the energy level of the site, located near the cytoplasmic channel (proton drain). In the first stage of this torque-generating process, the energy of the electrochemical potential is converted into potential energy of the proton-binding sites on the rotor. Afterwards, the tangential component of the Coulomb force produces a mechanical torque. We demonstrate that, at low temperatures, the loaded motor works in the shuttling regime where the energy of the electrochemical potential is consumed without producing any unidirectional rotation. The motor switches to the torque-generating regime at high temperatures, when the Brownian ratchet mechanism turns on. In the presence of a significant external torque, created by ATP hydrolysis, the system operates as a proton pump, which translocates protons against the transmembrane potential gradient. Here we focus on the F$_0$ motor, even though our analysis is applicable to the bacterial flagellar motor.Comment: 24 pages, 5 figure

Stiff Polymers, Foams and Fiber Networks

We study the elasticity of fibrous materials composed of generalized stiff polymers. It is shown that in contrast to cellular foam-like structures affine strain fields are generically unstable. Instead, a subtle interplay between the architecture of the network and the elastic properties of its building blocks leads to intriguing mechanical properties with intermediate asymptotic scaling regimes. We present exhaustive numerical studies based on a finite element method complemented by scaling arguments.Comment: 4 pages, 5 figure