Mechano-regulation by clathrin pit-formation and passive cholesterol-dependent tubules during de-adhesion.

Adherent cells ensure membrane homeostasis during de-adhesion by various mechanisms, including endocytosis. Although mechano-chemical feedbacks involved in this process have been studied, the step-by-step build-up and resolution of the mechanical changes by endocytosis are poorly understood. To investigate this, we studied the de-adhesion of HeLa cells using a combination of interference reflection microscopy, optical trapping and fluorescence experiments. We found that de-adhesion enhanced membrane height fluctuations of the basal membrane in the presence of an intact cortex. A reduction in the tether force was also noted at the apical side. However, membrane fluctuations reveal phases of an initial drop in effective tension followed by saturation. The area fractions of early (Rab5-labelled) and recycling (Rab4-labelled) endosomes, as well as transferrin-labelled pits close to the basal plasma membrane, also transiently increased. On blocking dynamin-dependent scission of endocytic pits, the regulation of fluctuations was not blocked, but knocking down AP2-dependent pit formation stopped the tension recovery. Interestingly, the regulation could not be suppressed by ATP or cholesterol depletion individually but was arrested by depleting both. The data strongly supports Clathrin and AP2-dependent pit-formation to be central to the reduction in fluctuations confirmed by super-resolution microscopy. Furthermore, we propose that cholesterol-dependent pits spontaneously regulate tension under ATP-depleted conditions.

Kolmogorov Turbulence Coexists with Pseudo-Turbulence in Buoyancy-Driven Bubbly Flows.

We investigate the spectral properties of buoyancy-driven bubbly flows. Using high-resolution numerical simulations and phenomenology of homogeneous turbulence, we identify the relevant energy transfer mechanisms. We find (a) at a high enough Galilei number (ratio of the buoyancy to viscous forces) the velocity power spectrum shows the Kolmogorov scaling with a power-law exponent -5/3 for the range of scales between the bubble diameter and the dissipation scale (η). (b) For scales smaller than η, the physics of pseudo-turbulence is recovered.

Rate of formation of caustics in heavy particles advected by turbulence.

The rate of collision and the relative velocities of the colliding particles in turbulent flows are a crucial part of several natural phenomena, e.g. rain formation in warm clouds and planetesimal formation in protoplanetary discs. The particles are often modelled as passive, but heavy and inertial. Within this model, large relative velocities emerge due to formation of singularities (caustics) of the gradient matrix of the velocities of the particles. Using extensive direct numerical simulations of heavy particles in both two (direct and inverse cascade) and three-dimensional turbulent flows, we calculate the rate of formation of caustics, [Formula: see text] as a function of the Stokes number ([Formula: see text]). The best approximation to our data is [Formula: see text], in the limit [Formula: see text] where [Formula: see text] is a non-universal constant. This article is part of the theme issue 'Scaling the turbulence edifice (part 2)'.

The breakdown of Darcy's law in a soft porous material.

We perform direct numerical simulations of the flow through a model of deformable porous medium. Our model is a two-dimensional hexagonal lattice, with defects, of soft elastic cylindrical pillars, with elastic shear modulus G, immersed in a liquid. We use a two-phase approach: the liquid phase is a viscous fluid and the solid phase is modeled as an incompressible viscoelastic material, whose complete nonlinear structural response is considered. We observe that the Darcy flux (q) is a nonlinear function - steeper than linear - of the pressure-difference (ΔP) across the medium. Furthermore, the flux is larger for a softer medium (smaller G). We construct a theory of this super-linear behavior by modelling the channels between the solid cylinders as elastic channels whose walls are made of material with a linear constitutive relation but can undergo large deformation. Our theory further predicts that the flow permeability is an universal function of ΔP/G, which is confirmed by the present simulations.

Rheology of Confined Non-Brownian Suspensions.

We study the rheology of confined suspensions of neutrally buoyant rigid monodisperse spheres in plane-Couette flow using direct numerical simulations. We find that if the width of the channel is a (small) integer multiple of the sphere diameter, the spheres self-organize into two-dimensional layers that slide on each other and the effective viscosity of the suspension is significantly reduced. Each two-dimensional layer is found to be structurally liquidlike but its dynamics is frozen in time.

Intermittency in the not-so-smooth elastic turbulence.

Elastic turbulence is the chaotic fluid motion resulting from elastic instabilities due to the addition of polymers in small concentrations at very small Reynolds ( Re ) numbers. Our direct numerical simulations show that elastic turbulence, though a low Re phenomenon, has more in common with classical, Newtonian turbulence than previously thought. In particular, we find power-law spectra for kinetic energy E(k) ~ k-4 and polymeric energy Ep(k) ~ k-3/2, independent of the Deborah (De) number. This is further supported by calculation of scale-by-scale energy budget which shows a balance between the viscous term and the polymeric term in the momentum equation. In real space, as expected, the velocity field is smooth, i.e., the velocity difference across a length scale r, δu ~ r but, crucially, with a non-trivial sub-leading contribution r3/2 which we extract by using the second difference of velocity. The structure functions of second difference of velocity up to order 6 show clear evidence of intermittency/multifractality. We provide additional evidence in support of this intermittent nature by calculating moments of rate of dissipation of kinetic energy averaged over a ball of radius r, εr, from which we compute the multifractal spectrum.

Shear thickening in non-Brownian suspensions: an excluded volume effect.

Shear thickening appears as an increase of the viscosity of a dense suspension with the shear rate, sometimes sudden and violent at high volume fraction. Its origin for noncolloidal suspension with non-negligible inertial effects is still debated. Here we consider a simple shear flow and demonstrate that fluid inertia causes a strong microstructure anisotropy that results in the formation of a shadow region with no relative flux of particles. We show that shear thickening at finite inertia can be explained as an increase of the effective volume fraction when considering the dynamically excluded volume due to these shadow regions.

Active buckling of pressurized spherical shells: Monte Carlo simulation.

We study the buckling of pressurized spherical shells by Monte Carlo simulations in which the detailed balance is explicitly broken-thereby driving the shell to be active, out of thermal equilibrium. Such a shell typically has either higher (active) or lower (sedate) fluctuations compared to one in thermal equilibrium depending on how the detailed balance is broken. We show that, for the same set of elastic parameters, a shell that is not buckled in thermal equilibrium can be buckled if turned active. Similarly a shell that is buckled in thermal equilibrium can unbuckle if sedated. Based on this result, we suggest that it is possible to experimentally design microscopic elastic shells whose buckling can be optically controlled.

Dynamic multiscaling in stochastically forced Burgers turbulence.

We carry out a detailed study of dynamic multiscaling in the turbulent nonequilibrium, but statistically steady, state of the stochastically forced one-dimensional Burgers equation. We introduce the concept of interval collapse time, which we define as the time taken for a spatial interval, demarcated by a pair of Lagrangian tracers, to collapse at a shock. By calculating the dynamic scaling exponents of the moments of various orders of these interval collapse times, we show that (a) there is not one but an infinity of characteristic time scales and (b) the probability distribution function of the interval collapse times is non-Gaussian and has a power-law tail. Our study is based on (a) a theoretical framework that allows us to obtain dynamic-multiscaling exponents analytically, (b) extensive direct numerical simulations, and (c) a careful comparison of the results of (a) and (b). We discuss possible generalizations of our work to higher dimensions, for the stochastically forced Burgers equation, and to other compressible flows that exhibit turbulence with shocks.

Large is different: Nonmonotonic behavior of elastic range scaling in polymeric turbulence at large Reynolds and Deborah numbers.

We use direct numerical simulations to study homogeneous and isotropic turbulent flows of dilute polymer solutions at high Reynolds and Deborah numbers. We find that for small wave numbers k, the kinetic energy spectrum shows Kolmogorov-like behavior that crosses over at a larger k to a novel, elastic scaling regime, E(k) ∼ k-ξ, with ξ ≈ 2.3. We study the contribution of the polymers to the flux of kinetic energy through scales and find that it can be decomposed into two parts: one increase in effective viscous dissipation and a purely elastic contribution that dominates over the nonlinear flux in the range of k over which the elastic scaling is observed. The multiscale balance between the two fluxes determines the crossover wave number that depends nonmonotically on the Deborah number. Consistently, structure functions also show two scaling ranges, with intermittency present in both of them in equal measure.

Chaos and irreversibility of a flexible filament in periodically driven Stokes flow.

The flow of Newtonian fluid at low Reynolds number is, in general, regular and time-reversible due to absence of nonlinear effects. For example, if the fluid is sheared by its boundary motion that is subsequently reversed, then all the fluid elements return to their initial positions. Consequently, mixing in microchannels happens solely due to molecular diffusion and is very slow. Here, we show, numerically, that the introduction of a single, freely floating, flexible filament in a time-periodic linear shear flow can break reversibility and give rise to chaos due to elastic nonlinearities, if the bending rigidity of the filament is within a carefully chosen range. Within this range, not only the shape of the filament is spatiotemporally chaotic, but also the flow is an efficient mixer. Overall, we find five dynamical phases: the shape of a stiff filament is time-invariant-either straight or buckled; it undergoes a period-two bifurcation as the filament is made softer; becomes spatiotemporally chaotic for even softer filaments but, surprisingly, the chaos is suppressed if bending rigidity is decreased further.

Dynamic multiscaling in two-dimensional fluid turbulence.

We obtain, by extensive direct numerical simulations, time-dependent and equal-time structure functions for the vorticity, in both quasi-Lagrangian and Eulerian frames, for the direct-cascade regime in two-dimensional fluid turbulence with air-drag-induced friction. We show that different ways of extracting time scales from these time-dependent structure functions lead to different dynamic-multiscaling exponents, which are related to equal-time multiscaling exponents by different classes of bridge relations; for a representative value of the friction we verify that, given our error bars, these bridge relations hold.

Persistence problem in two-dimensional fluid turbulence.

We present a natural framework for studying the persistence problem in two-dimensional fluid turbulence by using the Okubo-Weiss parameter Λ to distinguish between vortical and extensional regions. We then use a direct numerical simulation of the two-dimensional, incompressible Navier-Stokes equation with Ekman friction to study probability distribution functions (PDFs) of the persistence times of vortical and extensional regions by employing both Eulerian and Lagrangian measurements. We find that, in the Eulerian case, the persistence-time PDFs have exponential tails; by contrast, this PDF for Lagrangian particles, in vortical regions, has a power-law tail with an exponent θ=2.9±0.2.

Numerical study of large-scale vorticity generation in shear-flow turbulence.

Simulations of stochastically forced shear-flow turbulence in a shearing-periodic domain are used to study the spontaneous generation of large-scale flow patterns in the direction perpendicular to the plane of the shear. Based on an analysis of the resulting large-scale velocity correlations it is argued that the mechanism behind this phenomenon could be the mean-vorticity dynamo effect pioneered by Elperin, Kleeorin, and Rogachevskii [Phys. Rev. E 68, 016311 (2003)]. This effect is based on the anisotropy of the eddy viscosity tensor. One of its components may be able to replenish cross-stream mean flows by acting upon the streamwise component of the mean flow. Shear, in turn, closes the loop by acting upon the cross-stream mean flow to produce stronger streamwise mean flows. The diagonal component of the eddy viscosity is found to be of the order of the rms turbulent velocity divided by the wave number of the energy-carrying eddies.

Kazantsev dynamo in turbulent compressible flows.

We consider the kinematic fluctuation dynamo problem in a flow that is random, white-in-time, with both solenoidal and potential components. This model is a generalization of the well-studied Kazantsev model. If both the solenoidal and potential parts have the same scaling exponent, then, as the compressibility of the flow increases, the growth rate decreases but remains positive. If the scaling exponents for the solenoidal and potential parts differ, in particular if they correspond to typical Kolmogorov and Burgers values, we again find that an increase in compressibility slows down the growth rate but does not turn it off. The slow down is, however, weaker and the critical magnetic Reynolds number is lower than when both the solenoidal and potential components display the Kolmogorov scaling. Intriguingly, we find that there exist cases, when the potential part is smoother than the solenoidal part, for which an increase in compressibility increases the growth rate. We also find that the critical value of the scaling exponent above which a dynamo is seen is unity irrespective of the compressibility. Finally, we realize that the dimension d = 3 is special, as for all other values of d the critical exponent is higher and depends on the compressibility.

Clustering and energy spectra in two-dimensional dusty gas turbulence.

We present direct numerical simulation of heavy inertial particles (dust) immersed in two-dimensional turbulent flow (gas). The dust particles are modeled as monodispersed heavy particles capable of modifying the flow through two-way coupling. By varying the Stokes number (St) and the mass-loading parameter (ϕ_{m}), we study the clustering phenomenon and the gas phase kinetic energy spectra. We find that the dust-dust correlation dimension (d_{2}) also depends on ϕ_{m}. In particular, clustering decreases as mass loading (ϕ_{m}) is increased. In the kinetic energy spectra of gas we show (i) the emergence of a different scaling regime and that (ii) the scaling exponent in this regime is not universal but a function of both St and ϕ_{m}. Using a scale-by-scale enstrophy budget analysis we show that in this emerged scaling regime, which we call the dust-dissipative range, viscous dissipation due to the gas balances the back-reaction from the dust.

Statistics of the relative velocity of particles in turbulent flows: Monodisperse particles.

We use direct numerical simulations to calculate the joint probability density function of the relative distance R and relative radial velocity component V_{R} for a pair of heavy inertial particles suspended in homogeneous and isotropic turbulent flows. At small scales the distribution is scale invariant, with a scaling exponent that is related to the particle-particle correlation dimension in phase space, D_{2}. It was argued [K. Gustavsson and B. Mehlig, Phys. Rev. E 84, 045304 (2011)PLEEE81539-375510.1103/PhysRevE.84.045304; J. Turbul. 15, 34 (2014)1468-524810.1080/14685248.2013.875188] that the scale invariant part of the distribution has two asymptotic regimes: (1) |V_{R}|≪R, where the distribution depends solely on R, and (2) |V_{R}|≫R, where the distribution is a function of |V_{R}| alone. The probability distributions in these two regimes are matched along a straight line: |V_{R}|=z^{*}R. Our simulations confirm that this is indeed correct. We further obtain D_{2} and z^{*} as a function of the Stokes number, St. The former depends nonmonotonically on St with a minimum at about St≈0.7 and the latter has only a weak dependence on St.

Heavy inertial particles in turbulent flows gain energy slowly but lose it rapidly.

We present an extensive numerical study of the time irreversibility of the dynamics of heavy inertial particles in three-dimensional, statistically homogeneous, and isotropic turbulent flows. We show that the probability density function (PDF) of the increment, W(τ), of a particle's energy over a time scale τ is non-Gaussian, and skewed toward negative values. This implies that, on average, particles gain energy over a period of time that is longer than the duration over which they lose energy. We call this slow gain and fast loss. We find that the third moment of W(τ) scales as τ^{3} for small values of τ. We show that the PDF of power-input p is negatively skewed too; we use this skewness Ir as a measure of the time irreversibility and we demonstrate that it increases sharply with the Stokes number St for small St; this increase slows down at St≃1. Furthermore, we obtain the PDFs of t^{+} and t^{-}, the times over which p has, respectively, positive or negative signs, i.e., the particle gains or loses energy. We obtain from these PDFs a direct and natural quantification of the slow gain and fast loss of the energy of the particles, because these PDFs possess exponential tails from which we infer the characteristic loss and gain times t_{loss} and t_{gain}, respectively, and we obtain t_{loss}

How long do particles spend in vortical regions in turbulent flows?

We obtain the probability distribution functions (PDFs) of the time that a Lagrangian tracer or a heavy inertial particle spends in vortical or strain-dominated regions of a turbulent flow, by carrying out direct numerical simulations of such particles advected by statistically steady, homogeneous, and isotropic turbulence in the forced, three-dimensional, incompressible Navier-Stokes equation. We use the two invariants, Q and R, of the velocity-gradient tensor to distinguish between vortical and strain-dominated regions of the flow and partition the Q-R plane into four different regions depending on the topology of the flow; out of these four regions two correspond to vorticity-dominated regions of the flow and two correspond to strain-dominated ones. We obtain Q and R along the trajectories of tracers and heavy inertial particles and find out the time t_{pers} for which they remain in one of the four regions of the Q-R plane. We find that the PDFs of t_{pers} display exponentially decaying tails for all four regions for tracers and heavy inertial particles. From these PDFs we extract characteristic time scales, which help us to quantify the time that such particles spend in vortical or strain-dominated regions of the flow.

Deviation-angle and trajectory statistics for inertial particles in turbulence.

Small particles in suspension in a turbulent fluid have trajectories that do not follow the pathlines of the flow exactly. We investigate the statistics of the angle of deviation ϕ between the particle and fluid velocities. We show that, when the effects of particle inertia are small, the probability distribution function (PDF) P_{ϕ} of this deviation angle shows a power-law region in which P_{ϕ}∼ϕ^{-4}. We also find that the PDFs of the trajectory curvature κ and modulus θ of the torsion ϑ have power-law tails that scale, respectively, as P_{κ}∼κ^{-5/2}, as κ→∞, and P_{θ}∼θ^{-3}, as θ→∞: These exponents are in agreement with those previously observed for fluid pathlines. We propose a way to measure the complexity of heavy-particle trajectories by the number N_{I}(t,St) of points (up until time t) at which the torsion changes sign. We present numerical evidence that n_{I}(St)≡lim_{t→∞}N_{I}(t,St)/t∼St^{-Δ} for large St, with Δ≃0.5.