Budding and Domain Shape Transformations in Mixed Lipid Films and Bilayer Membranes

We study the stability and shapes of domains with spontaneous curvature in fluid films and membranes, embedded in a surrounding membrane with zero spontaneous curvature. These domains can result from the inclusion of an impurity in a fluid membrane, or from phase separation within the membrane. We show that for small but finite line and surface tensions and for finite spontaneous curvatures, an equilibrium phase of protruding circular domains is obtained at low impurity concentrations. At higher concentrations, we predict a transition from circular domains, or "caplets", to stripes. In both cases, we calculate the shapes of these domains within the Monge representation for the membrane shape. With increasing line tension, we show numerically that there is a budding transformation from stable protruding circular domains to spherical buds. We calculate the full phase diagram, and demonstrate a two triple points, of respectively bud-flat-caplet and flat-stripe-caplet coexistence.Comment: 14 pages, to appear in Phys Rev

Poisson's ratio in composite elastic media with rigid rods

We study the elastic response of composites of rods embedded in elastic media. We calculate the micro-mechanical response functions, and bulk elastic constants as functions of rod density. We find two fixed points for Poisson's ratio with respect to the addition of rods in 3D composites: there is an unstable fixed point for Poisson's ratio=1/2 (an incompressible system) and a stable fixed point for Poisson's ratio=1/4 (a compressible system). We also derive an approximate expression for the elastic constants for arbitrary rod density that yields exact results for both low and high density. These results may help to explain recent experiments [Physical Review Letters 102, 188303 (2009)] that reported compressibility for composites of microtubules in F-actin networks.Comment: 4 pages, 4 figures, to appear in Phys. Rev. Let

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

Inspiration for the Future: The Role of Inspiratory Muscle Training in Cystic Fibrosis.

Cystic fibrosis (CF) is an inherited, multi-system, life-limiting disease characterized by a progressive decline in lung function, which accounts for the majority of CF-related morbidity and mortality. Inspiratory muscle training (IMT) has been proposed as a rehabilitative strategy to treat respiratory impairments associated with CF. However, despite evidence of therapeutic benefits in healthy and other clinical populations, the routine application of IMT in CF can neither be supported nor refuted due to the paucity of methodologically rigorous research. Specifically, the interpretation of available studies regarding the efficacy of IMT in CF is hampered by methodological threats to internal and external validity. As such, it is important to highlight the inherent risk of bias that differences in patient characteristics, IMT protocols, and outcome measurements present when synthesizing this literature prior to making final clinical judgments. Future studies are required to identify the characteristics of individuals who may respond to IMT and determine whether the controlled application of IMT can elicit meaningful improvements in physiological and patient-centered clinical outcomes. Given the equivocal evidence regarding its efficacy, IMT should be utilized on a case-by-case basis with sound clinical reasoning, rather than simply dismissed, until a rigorous evidence-based consensus has been reached

The mechanical response of semiflexible networks to localized perturbations

Previous research on semiflexible polymers including cytoskeletal networks in cells has suggested the existence of distinct regimes of elastic response, in which the strain field is either uniform (affine) or non-uniform (non-affine) under external stress. Associated with these regimes, it has been further suggested that a new fundamental length scale emerges, which characterizes the scale for the crossover from non-affine to affine deformations. Here, we extend these studies by probing the response to localized forces and force dipoles. We show that the previously identified nonaffinity length [D.A. Head et al. PRE 68, 061907 (2003).] controls the mesoscopic response to point forces and the crossover to continuum elastic behavior at large distances.Comment: 16 pages, 18 figures; substantial changes to text and figures to clarify the crossover to continuum elasticity and the role of finite-size effect

Non-equilibrium microtubule fluctuations in a model cytoskeleton

Biological activity gives rise to non-equilibrium fluctuations in the cytoplasm of cells; however, there are few methods to directly measure these fluctuations. Using a reconstituted actin cytoskeleton, we show that the bending dynamics of embedded microtubules can be used to probe local stress fluctuations. We add myosin motors that drive the network out of equilibrium, resulting in an increased amplitude and modified time-dependence of microtubule bending fluctuations. We show that this behavior results from step-like forces on the order of 10 pN driven by collective motor dynamics

One-Bead Microrheology with Rotating Particles

We lay the theoretical basis for one-bead microrheology with rotating particles, i.e, a method where colloids are used to probe the mechanical properties of viscoelastic media. Based on a two-fluid model, we calculate the compliance and discuss it for two cases. We first assume that the elastic and fluid component exhibit both stick boundary conditions at the particle surface. Then, the compliance fulfills a generalized Stokes law with a complex shear modulus whose validity is only limited by inertial effects, in contrast to translational motion. Secondly, we find that the validity of the Stokes regime is reduced when the elastic network is not coupled to the particleComment: 7 pages, 5 figures, submitted to Europhys. Let

Short-time inertial response of viscoelastic fluids measured with Brownian motion and with active probes

We have directly observed short-time stress propagation in viscoelastic fluids using two optically trapped particles and a fast interferometric particle-tracking technique. We have done this both by recording correlations in the thermal motion of the particles and by measuring the response of one particle to the actively oscillated second particle. Both methods detect the vortex-like flow patterns associated with stress propagation in fluids. This inertial vortex flow propagates diffusively for simple liquids, while for viscoelastic solutions the pattern spreads super-diffusively, dependent on the shear modulus of the medium

One- and two-particle microrheology

We study the dynamics of rigid spheres embedded in viscoelastic media and address two questions of importance to microrheology. First we calculate the complete response to an external force of a single bead in a homogeneous elastic network viscously coupled to an incompressible fluid. From this response function we find the frequency range where the standard assumptions of microrheology are valid. Second we study fluctuations when embedded spheres perturb the media around them and show that mutual fluctuations of two separated spheres provide a more accurate determination of the complex shear modulus than do the fluctuations of a single sphere.Comment: 4 pages, 1 figur