Anomalous transport in the crowded world of biological cells

A ubiquitous observation in cell biology is that diffusion of macromolecules and organelles is anomalous, and a description simply based on the conventional diffusion equation with diffusion constants measured in dilute solution fails. This is commonly attributed to macromolecular crowding in the interior of cells and in cellular membranes, summarising their densely packed and heterogeneous structures. The most familiar phenomenon is a power-law increase of the MSD, but there are other manifestations like strongly reduced and time-dependent diffusion coefficients, persistent correlations, non-gaussian distributions of the displacements, heterogeneous diffusion, and immobile particles. After a general introduction to the statistical description of slow, anomalous transport, we summarise some widely used theoretical models: gaussian models like FBM and Langevin equations for visco-elastic media, the CTRW model, and the Lorentz model describing obstructed transport in a heterogeneous environment. Emphasis is put on the spatio-temporal properties of the transport in terms of 2-point correlation functions, dynamic scaling behaviour, and how the models are distinguished by their propagators even for identical MSDs. Then, we review the theory underlying common experimental techniques in the presence of anomalous transport: single-particle tracking, FCS, and FRAP. We report on the large body of recent experimental evidence for anomalous transport in crowded biological media: in cyto- and nucleoplasm as well as in cellular membranes, complemented by in vitro experiments where model systems mimic physiological crowding conditions. Finally, computer simulations play an important role in testing the theoretical models and corroborating the experimental findings. The review is completed by a synthesis of the theoretical and experimental progress identifying open questions for future investigation.Comment: review article, to appear in Rep. Prog. Phy

Structural relaxation in orthoterphenyl: a schematic mode coupling theory model analysis

Depolarized light scattering spectra of orthoterphenyl showing the emergence of the structural relaxation below the oscillatory microscopic excitations are described by solutions of a schematic mode--coupling--theory model

Shear response of a smectic film stabilized by an external field

The response of a field-stabilized two-dimensional smectic to shear stress is discussed. Below a critical temperature the smectic film exhibits elastic response to an infinitesimal shear stress normal to the layering. At finite stresses free dislocations nucleate and relax the applied stress. The coupling of the dislocation current to the stress results in non-newtonian viscous flow. The flow profile in a channel geometry is shown to change qualitatively from a power-law dependence to a Poiseuille-like profile opon increasing the pressure head

How Glassy Relaxation Slows Down by Increasing Mobility

We investigate how structural relaxation in mixtures with strong dynamical asymmetry is affected by the microscopic dynamics. Brownian and Newtonian dynamics simulations of dense mixtures of fast and slow hard spheres reveal a striking trend reversal. Below a critical density, increasing the mobility of the fast particles fluidizes the system, yet, above that critical density, the same increase in mobility strongly hinders the relaxation of the slow particles. The critical density itself does not depend on the dynamical asymmetry and can be identified with the glass-transition density of the mode-coupling theory. The asymptotic dynamics close to the critical density is universal, but strong pre-asymptotic effects prevail in mixtures with additional size asymmetry. This observation reconciles earlier findings of a strong dependence on kinetic parameters of glassy dynamics in colloid--polymer mixtures with the paradigm that the glass transition is determined by the properties of configuration space alone

Intermediate scattering function of an anisotropic active Brownian particle

Various challenges are faced when animalcules such as bacteria, protozoa, algae, or sperms move autonomously in aqueous media at low Reynolds number. These active agents are subject to strong stochastic fluctuations, that compete with the directed motion. So far most studies consider the lowest order moments of the displacements only, while more general spatio-temporal information on the stochastic motion is provided in scattering experiments. Here we derive analytically exact expressions for the directly measurable intermediate scattering function for a mesoscopic model of a single, anisotropic active Brownian particle in three dimensions. The mean-square displacement and the non-Gaussian parameter of the stochastic process are obtained as derivatives of the intermediate scattering function. These display different temporal regimes dominated by effective diffusion and directed motion due to the interplay of translational and rotational diffusion which is rationalized within the theory. The most prominent feature of the intermediate scattering function is an oscillatory behavior at intermediate wavenumbers reflecting the persistent swimming motion, whereas at small length scales bare translational and at large length scales an enhanced effective diffusion emerges. We anticipate that our characterization of the motion of active agents will serve as a reference for more realistic models and experimental observations.Comment: 10 pages, 4 figure

Driven lattice gas of dimers coupled to a bulk reservoir

We investigate the non-equilibrium steady state of a one-dimensional (1D) lattice gas of dimers. The dynamics is described by a totally asymmetric exclusion process (TASEP) supplemented by attachment and detachment processes, mimicking chemical equilibrium of the 1D driven transport with the bulk reservoir. The steady-state phase diagram, current and density profiles are calculated using both a refined mean-field theory and extensive stochastic simulations. As a consequence of the on-off kinetics, a new phase coexistence region arises intervening between low and high density phases such that the discontinuous transition line of the TASEP splits into two continuous ones. The results of the mean-field theory and simulations are found to coincide. We show that the physical picture obtained in the corresponding lattice gas model with monomers is robust, in the sense that the phase diagram changes quantitatively, but the topology remains unaltered. The mechanism for phase separation is identified as generic for a wide class of driven 1D lattice gases.Comment: 15 pages, 10 figures, 1tabl