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Swimming at small scales: mechanisms and emergent structures I. Pagonabarraga University of Barcelona 1. Introduction 2. Swimming at small scales 3. Model swimmers 4. Collective swimming 5. Chemical swimmers 6. Molecular motors 2 1. Introduction Active systems: collections of elements able to convert internal energy into mechanical work (autonomous motion) intrinsically out-of-equilibrium systems (even in a steady state, if any, and without external forcing) Examples flocks of birds bacterial colonies artificial self-propelled objects molecular motors 1. Introduction Flagella E.coli Volvox Amoeba Cilia Ciliated Microorganisms Paramecium 1. Introduction Actuated colloids Adding reactivity: new propelling mechanisms Different sets of micro/nano robots Heterogeneous particles Confinement + asymmetric mobility no deformation Desired structures, adaptive, capable of self repair 1. Introduction Energy from small scales Systems intrinsically out of equilibrium Nonequilibrium distributions no detailed balance Ratchets Emerging patterns and phases 2. Swimming Re = uL/ν 2. Swimming Re = uL/ν L~µm u~10 µm/s ν~10-2 cm2/s Inertia irrelevant Reciprocal motion (1 degree of freedom) Purcell, Am.J.Phys. (1977) Simple models: wave propagation body deformation Najafi et al Purcells’ theorem Reversibility: no net displacement Bibette et al 2. Swimming Re = uL/ν L~µm u~10 µm/s ν~10-2 cm2/s Inertia irrelevant Reciprocal motion (1 degree of freedom) Purcell, Am.J.Phys. (1977) Purcells’ theorem Reversibility: no net displacement Bibette et al 3. Model swimmers Each move change in HI Decompose cycle in 4 moves Total displacement in a cycle Mean velocity Expansion in small displacement 3. Model swimmers Each move change in HI Decompose cycle in 4 moves Low Reynolds Hydrodynamic coupling Oseen tensor Force free motion 3. Actuated swimmers Magnetic field: Local acceleration Asymmetry + local bending: wave Top view Doublet vs isolated Bibette et al Confinement + asymmetric mobility no deformation Larger aggregate 3. Actuated swimmers Difference in friction along cycle Tierno et al. PRL 2010 Open trajectories: no net motion Persistent ballistic motion 3. Actuated swimmers Two regimes: optimal velocity Different sensitivity to driving General behavior for general assemblies Paramagnetic colloids 3. Actuated swimmers Pair velocities: Low Re Constraint: COM + orientation linear relation Effective mobilities Asymmetric particle/wall friction For circular trajectory: Net average velocity 3. Actuated swimmers Angular velocity: Low Re Kinematic constraint Circularly polarized magnetic field Characteristic frequency 3. Actuated swimmers Good agreement Fitting: height rotational friction Low Ω : increase rectification rate change in orientation negligible High Ω : doublet aligns parallel to wall decrease in asymmetry Controlled motion in microfluidic devices 4. Cilliated swimmers Squirmers Metachronal wave on Opalina, Paramecium. Fixed tangential velocity profile on the surface (Lighthill, 1952; Blake, 1971) Opalina Surface tangential velocity β=B2/B1 Propulsion velocity 4. Cilliated swimmers Squirmers Metachronal wave on Opalina, Paramecium. Fixed tangential velocity profile on the Opalina surface (Lighthill, 1952; Blake, 1971) Balantidium coli Surface tangential velocity oligotrich β=B2/B1 Propulsion velocity 4. Cilliated swimmers Collective distorsion of cillia small compared with body object effective waves Time-dependent slip velocity Solve Stokes equation 4. Cilliated swimmers Force-free swimmer sets the self-propelling velocity Disregard normal oscillations + focus in average flow field Active stress induced by the effective slip velocity? 4. Cilliated swimmers Dynamics of active particles Low Reynolds numbers Absence of external driving closer to electrophoresis? Relevance of swimming mechanism Fluid flows with vorticity Coupling translation/rotation relevance of near field interactions chlamydomonas v~1/r2 β>0 Contractile β<0 Extensile B1=0 B2≠0 β=0 Passive squirmer Apolar v~1/r3 5. Squirmer suspensions Time scales: analogous to passive colloids τr ∼ R2/ν τr << τm << τD τm∼R/u∞ Diffusion induced by collisions τD∼R2/D Size~ 1-500 µm Speeds < 100 µm/s Neglect thermal fluctuations fluid inertia (Re<0.01) τD/τm = u∞ R/D= Pe > 1 Paramecium Opalina u∞ ~1000 µm/s u∞ ~100 µm/s R ~200 µm Re ~ 0.2 R ~200 µm Re ~ 0.02 5. Squirmer suspensions Intermediate times t ~ τm HI align propellers effective interactions Relevance near field couplings Particle acceleration Decay vacf Transient aggregate formation Pedley 2006 5. Squirmer suspensions Long times t ~ τD φ=0.1 Relevance near field couplings Particle acceleration φ=0.3 Structure: aggregates orientational order finite life time 5. Squirmer suspensions Mean square displacement 2D Super-diffusive regime Asymetric driving Low volume fractions φ=0.1 φ=0.4 Flocking models transitions at high φ Percolating clusters Additional interactions stabilize clusters Detailed analysis analogies with spps’ transitions 5. Squirmer suspensions Contractile 28 6.Colloidal phoresis Phoretic transport: motion of colloidal particle under the effect of external field (electric field, concentration/temperature gradients) (solute molecules size)/(colloidal particle size) large scale aggregates R (Sen et al, Faraday Discuss. 143, 15 (2009)) 6.Colloidal phoresis Concentration interacts with a solid surface delocalized membrane? Concentration gradient parallel to the surface asymmetric change in chemical potential Generates pressure gradient Due to asymmetry induced by wall Surface-induced flow 30 6. Colloidal phoresis The presence of the solute can be taken into account by means of an effective “slip velocity” as boundary condition for the flow on the colloid surface surface phoretic mobility colloid/solute interaction potential (short-ranged) velocity of a (spherical) particle of radius R (Anderson, ARFM 21, 61 (1985); Golestanian et al, New J. Phys. 9, 126 (2007)) Test case: chemotaxis Directed motion of a colloidal particle in a linear concentration profile For constant mobility the propulsion velocity can be calculated exactly chemoattractant chemorepellent Autonomous motion What if particles might generate concentration gradients? self-propulsion!!! (Paxton et al, JACS 126, 13424 (2004)) Janus (Golestanian et al, PRL 94, 22081 (2005)) activity modelled by simple updating rule non-conserved dynamics for φ! particle surface activity Janus particle The velocity of an isolated particle of constant mobility µ can be computed exactly 7.Collective dynamics in “2D”: phoretic mobility? 7. Density fluctuations 7. Repulsive chemical swimmers No hydrodynamics Towards a crystal structure Faster dynamics larger number of “defects” 7. Atractive chemical swimmers No hydrodynamics Cluster formation 37 7. Radial distribution functions 7. Statistics and geometry of clusters 7. Orientational radial distribution functions where being the angle between the orientation vector of the reference particle and the one of the generic particle in Hydrodynamics: induces particle rotation Leads to disorder / induced flows 10. Conclusions Active matter Release energy at small scales (natural/synthetic) intrinsically out of equilibrium New mechanisms to develop patterns and structures Variety of mechanisms to swim Mesoscopic modeling simple swimmers / chemical swimmers Identify chemical/hydrodynamic interactions Induced flows Cluster formation Enhanced fluctuations / pumping Phoretic mobility relevant role (non-eq) transition from repulsive crystal to clustering Chemical signaling mainly determines clustering hydrodynamics limits cluster growth