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Emergence of coherent structuresand large-scale flows in biologically
active suspensionsDavid Saintillan
Department of Mechanical Science and EngineeringUniversity of Illinois at Urbana-Champaign
IMA workshop: Natural locomotion in
fluids and on surfaces,
Minneapolis, MN, June 3 2010
Collaborator: Michael J. Shelley(Courant Institute, NYU)
Graduate student: Amir AlizadehPahlavan (MechSE, UIUC)
Interactions in active suspensions
E. Coli swarming on a surface
Active suspension: suspensions of self-propelled particles (e.g. bacteria, microalgae,artificial microswimmers).
Experiments exhibit:! correlated motions over length scales thatgreatly exceed the particle dimensions.! diffusive behavior! in some cases, large-scale densityfluctuations suggesting self-organization withinthe suspensions.Dombrowski et al., PRL (2004)Mendelson et al., J. Bacteriol. (1999)Wu and Libchaber, PRL (2000)Soni et al., Biophys. J. (2003)Sokolov et al., PRL (2007)and many others…
! What mechanisms lead to these large-scaleflows and pattern formation?! Under which conditions (concentration,system size, swimming mechanism) do theseoccur?! What are the characteristics of the resultingflows and patterns?
P. Mirabilis swarming on a surface
(Weibel Lab, University of Wisconsin)
Swimming at low Reynolds numberIn the absence of inertia, swimming must rely of non-reciprocal shape deformations. Innature, several mechanisms for swimming are observed:! flagellar propulsion (sperm, E Coli, B Subtilis…)! ciliary propulsion (Paramecium…)! body deformation (C Elegans…)
sperm Chlamydomonas Paramecium C Elegans
“Life at low Reynolds number”E.M. Purcell 1977
pusher particle
In all cases, the swimming particle exerts a propulsive force on the fluid, which isexactly balanced by the viscous drag: to leading order, it creates a force dipole.
puller particle
Kinetic theory: governing equations
! Continuity equation for the distribution function of particle center-of-mass and director :
! Flux velocities:
! Fluid disturbance velocity:
Particle extra stress:
Stress magnitude and swimming velocity arerelated on the particle scale by:
! Non-dimensionalization:
Saintillan & Shelley PRL 100 178103 (2008), Phys. Fluids 20 123304 (2008)Hohenegger & Shelley, PRE 81 046311 (2010)
Configurational entropy
! We define the system entropy as:
for an isotropic homogeneous suspension
! The evolution of the entropy is governed by the following equation:
diffusive damping
Pullers : the entropy is driven down by both diffusive processes and viscous dissipation
Pushers : there is a feedback loop wherein fluctuations create velocity gradients which increase fluctuations
Reduced equations for aligned case
! Consider a locally aligned suspension:
Particle extra stress:
! If diffusion can be neglected, the full kinetic equations reduce to the followingequations for the concentration field and director field:
! The disturbance velocity field then satisfies:
Stability analysis: aligned case
! Consider a nearly aligneduniform suspension:
! The two growth rates have oppositesigns: the suspension is alwaysunstable.
! In the long wave limit, the solution ofSimha and Ramaswamy (2002) isrecovered:
! In real systems, diffusion will damphigh-wavenumber fluctuations.
Stability analysis: uniform isotropic case
! Nearly isotropic uniform suspension:
Eigenvalue problem for particle stress tensor:
Eigenvectors are of the form:
Dispersion relation:
Suspensions ofpushers areunstable for
For more details on this problem see Christel Hohenegger’s talk on Saturday.
Nonlinear simulations
Integrate the kineticequations using
! a spectral solution of theStokes equations
! 2nd-order finitedifferences forconservation equation
! 2nd order Adams-Bashforth time marching.
Parallel code, typicallyrun on 64 processors.
Concentration fluctuations
Concentration autocorrelation functionConcentration isosurfaces (c = 1.5)
! Concentration fluctuations are in the form of transient sheets that constantly formand break up in time in a quasi-periodic fashion.
! The concentration field is correlated on large-length scales, with anautocorrelation length of the order of 1/4 of the box size.
Disturbance velocity field
! Velocity fields are chaotic, correlated onscales of the order of the system size, andstrongly dominated by large-scales (rapiddecay of velocity power spectrum withwavenumber)
-4.5
Velocity power spectrum
Velocity autocorrelation function
Evolution of Fourier modes
! Decay of high-wavenumber fluctuations with oscillations
! Growth of low-wavenumber fluctuations. At long-times, quasi-periodic oscillationsare observed over long time scales.
! Shear stresses and director divergence grow first, followed by concentration.
Mechanism for concentration fluctuations
Evolution equation for concentration field:
source term
Mechanism for the formation of concentration fluctuations:
! The stress instability causes the particles to align locally (particles stress = orderparameter)
! Particles swim towards regions of negative
Entropy and power input
Note:
power input rate of viscousdissipation
pushers pushers
pullers pullers
Slender-body theory for swimming rod
tangentialtractions
normaltractions
! Surface tractions
! Integrated traction (force per unit length):
! Force and torque balances
Saintillan & Shelley, Phys. Rev. Lett. 99 058102 (2007)
Slender-body theory for swimming rod
prescribed unknownSlip velocity(unknown)
Disturbancefluid velocity
Green’s function for Stokes flow
unknown
Local slender-body equation
! with prescribed stress:
! with no-slip boundary condition:
Disturbance fluid velocity (hydrodynamic interactions):
! Coupled linear system of integral equations, solved by spectral methods:
- Legendre polynomial expansion of the force distribution - Smooth particle mesh algorithm (Saintillan, Darve & Shaqfeh, Phys. Fluids 2005)
Batchelor JFM (1971)
Particle dynamicsPushers
800 particles
Pushers
4000 particles
Pushers
8000 particles
Pullers
8000 particles
Disturbance velocity field
Onset of collectiveswimming
Pushers Pushers
Pushers Pullers
Large-scale correlated flows appearin suspensions of pushers witheffective volume fraction above 0.5.
Number density fluctuations! Particle occupancy statistics: statistics of the number of particles falling inside acubic interrogation cell placed at an arbitrary location inside the simulation box.
! Poisson law for a random distribution of particles:
Number density fluctuations
Onset ofcollectiveswimming
! Standard deviation of number density fluctuations exhibitspower law dependence on the mean number of particles:
! For a Poisson law,
Orientation correlations
Orientationally correlated regions
Pair correlation function for orientations
Velocities and local structure
High particle velocities correlatewith:
! high local density (clusters)
! local polar alignment
Orientational diffusion
Define rotary diffusion coefficientas the inverse time scale fororientation decorrelation:
Rotary diffusion coefficient
Linear increase of rotary diffusion coefficient at low concentration as a result ofpair interactions.
Translational diffusionIsotropic translational diffusion coefficientTwo definitions for translational
diffusion coefficient:
Decrease of translational diffusion coefficient with concentration: diffusion arisesthrough randomization of orientation by angular diffusion (generalized Taylordispersion process, cf. Brenner 1980).
Fluid mixingPushers
800 particles
Pushers
4000 particles
Pullers
8000 particles
Pushers
8000 particles
Fluid mixing
Onset ofcollectiveswimming
Multiscale mixing norm:
(Mathew, Mezic & Petzold, Physica D 211 23-46 2005)
mixing efficiency b
Rheology of swimming suspensions
Recent experiments show a decrease in viscosity in suspensions of swimmingB. Subtilis (bacterium, pusher)
Bacterial concentration Bacterial mean swimming speed
Sokolov & Aranson, “Reduction of viscosity in a suspension ofswimming bacteria”, PRL 103, 148101 (2009)
Rheology of swimming suspensions
In suspensions of Chlamydomonas (alga, puller), however, a viscosityenhancement is observed.
Rafai, Jibuti & Peyla, “Effective viscosity of microswimmersuspensions”, Phys. Rev. Lett. 104 098102 (2010)
Dilute rheology: Kinetic theory (1/3)
! Interparticle hydrodynamic interactionsare neglected.
! We consider a single particle swimmingin an imposed shear flow at low Reynoldsnumber.
! The configuration of the suspension isentirely determined by the orientationdistribution .
! Fokker-Planck equation for the orientation distribution:
shear flow diffusion tumbling
! Rotational velocity (Jeffery 1922):
(Subramanian andKoch JFM 2009)
Saintillan, Exp. Mech. (to appear, 2010), Phys. Rev. E 81 056307 (2010)
Dilute rheology: Kinetic theory (2/3)! Stress calculation: ensemble average of the force dipoles on the particles
Newtonian stress particle extra stress
Total stress:
Batchelor (Annu. Rev. Fluid Mech. 1974), Hinch & Leal (JFM 1972, 1975)
Particle extra stress:
Three contributions: External flow
“Brownian” rotations
Dipole due to swimming
For a slender-body:(Hinch & Leal, JFM 1976)
Pusher: Puller:
Dilute rheology: Kinetic theory (3/3)! Characteristic scales and non-dimensionalization
Particle viscosity
First and secondnormal stressdifferencecoefficients
! Dimensionless equations
Characteristic scales
Dimensionless parameters
Steady-state FP equation
Particle extra stress
Pusher:
Puller:
Orientation distributions
! The steady-state Fokker-Planckequation is solved spectrally byexpanding the orientationdistribution on the basis ofspherical harmonics.
! Numerical solutions wereobtained using harmonics ofdegree up to 40 (corresponding to861 modes), which ensured anexcellent accuracy for theparameters considered here.
Chen & Koch (Phys. Fluids 1996)Doi & Edwards (J. Chem. Soc. 1978)
Negative particle viscosity
! Zero-shear-rate particle viscosity ( ):
! Negative particle viscosities occur for: ( for slender swimmers)
! Critical shear ratebelow which negativeviscosities occur vs.rotary diffusivity
Effect of particle tumbling
Slender swimmerswith fixed rotary diffusivity
Particle tumbling has the samequalitative effect as rotary diffusion.
Extensional rheology
! The same model can be developed for uniaxial extensional and compressional flows,as well as planar extensional flows, and is amenable to analytical solutions.
Saintillan, Phys. Rev. E 81 056307 (2010)
Stability in shear flow
Substituting in the governing equations, the linearized equation becomes:
This is a generalized eigenvalue problem (integro-differential equation) which can beturned into a system of algebraic eigenvalue problems using spherical harmonics.
Saintillan and AlizadehPahlavan, in preparation 2010
Summary! A kinetic theory for suspensions of self-propelled microswimmers predicts aninstability in isotropic suspensions of pushers, which occurs above a certainsystem size and/or suspension volume fraction. The instability is characterized bythe growth of density fluctuations, in the form of dense clusters that form andbreak up in time.
! Particle simulations confirm the results of the kinetic model, and show theemergence of coherent structures and large-scale flows in concentratedsuspensions of pushers, as demonstrated by an increase in density fluctuations,velocity correlation length, and mean particle velocity above a threshold volumefraction. These effects are not observed for pullers.
! The large-scale flows that result from the instability cause very efficient fluidmixing.
! The effective rheology of active suspensions is characterized by a decrease inviscosity in weak flows of pushers, and by an enhancement for pullers.
! An external shear flow has a stabilizing effect. Kinetic simulations show atransition from 3D to 2D to 1D instability as shear rate increases, followed bycomplete stabilization.
! Acknowledgments: NSF Grant DMS-0920931-ARRA, National Center forSupercomputing Applications